diversity and palaeoecology of early devonian invertebrate ... · of morocco (e.g. klug et al....

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Year: 2014

Diversity and palaeoecology of Early Devonian invertebrate associations inthe Tafilalt (Anti-Atlas, Morocco)

Frey, L <javascript:contributorCitation( ’Frey, L’ );>; Naglic, C <javascript:contributorCitation(’Naglic, C’ );>; Hofmann, R <javascript:contributorCitation( ’Hofmann, R’ );>; Schemm-Gregory, M<javascript:contributorCitation( ’Schemm-Gregory, M’ );>; Fryda, J <javascript:contributorCitation(

’Fryda, J’ );>; Kröger, B <javascript:contributorCitation( ’Kröger, B’ );>; Taylor, P D<javascript:contributorCitation( ’Taylor, P D’ );>; Wilson, M A <javascript:contributorCitation(

’Wilson, M A’ );>; Klug, C <javascript:contributorCitation( ’Klug, C’ );>

DOI: https://doi.org/10.3140/bull.qeosci.1459

Posted at the Zurich Open Repository and Archive, University of ZurichZORA URL: https://doi.org/10.5167/uzh-85791Journal ArticlePublished Version

Originally published at:Frey, L; Naglic, C; Hofmann, R; Schemm-Gregory, M; Fryda, J; Kröger, B; Taylor, P D; Wilson, M A;Klug, C (2014). Diversity and palaeoecology of Early Devonian invertebrate associations in the Tafilalt(Anti-Atlas, Morocco). Bulletin of Geosciences, 89(1):75-112.DOI: https://doi.org/10.3140/bull.qeosci.1459

https://doi.org/10.3140/bull.qeosci.1459

https://doi.org/10.5167/uzh-85791

https://doi.org/10.3140/bull.qeosci.1459

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Quantitative analyses of the taxonomic composition and palaeoecology of five Early Devonian faunules (earliestLochkovian to early Emsian) collected from the locality Jebel Ouaoufilal in the Tafilalt (Morocco) were conducted. Weexamined 3376 specimens belonging to 158 species and their stratigraphic distribution. The quantitative data sets wereanalysed for alpha diversity and ecospace utilization. Macrofossils of every faunule were identified, counted andgrouped according to ecological categories of tiering, motility and feeding behaviour. Based on these data, we noted (i) astrong increase in species richness, especially in benthic species, from the lowermost Lochkovian to the Pragian stageand a following subtle decrease in the early Emsian and (ii) a considerable expansion of ecospace use from the earliestLochkovian to the early Emsian. These general trends are considered to be a result of favourable living conditions due toa growing oxygen content at the sea floor, which correlates with a regional and potentially a global regression from theLochkovian to the Pragian. A transgression in the early Zlíchovian probably reduced the species richness by decreasingthe oxygenation of bottom waters. Additionally, we described and figured three new genera and six new species of thediverse Pragian faunule (86 species): the hederelloid Filihernodia buccina gen. et sp. nov., the crinoid Hexawacrinusclaudiakurtae gen. et sp. nov., the gastropods Oriomphalus multiornatus sp. nov., Eohormotomina restisevoluta gen. etsp. nov., and the cephalopods Tenuitheoceras secretum gen. et sp. nov. and Arionoceras kennethdebaetsi sp. nov. We re-cord the first occurrence of Tiaracrinus moravicus from Africa. • Key words: Early Devonian, alpha diversity, palaeoec-ology, Cephalopoda, Gastropoda, Anti-Atlas, Morocco.

FREY, L., NAGLIK, C., HOFMANN, R., SCHEMM-GREGORY, M., FRÝDA, J., KRÖGER, B., TAYLOR, P.D., WILSON, M.A.& KLUG, C. 2014. Diversity and palaeoecology of Early Devonian invertebrate associations in the Tafilalt (Anti-Atlas,Morocco). Bulletin of Geoscience 89(1), 75–112 (12 figures, appendix). Czech Geological Survey, Prague. ISSN1214-1119. Manuscript received June 13, 2013; accepted in revised form October 29, 2013; published online December12, 2013; issued xxxxxxx xx, 2013.

Linda Frey, Carole Naglik, Richard Hofmann & Christian Klug, Palaeontological Institute and Museum, University ofZurich, Karl Schmid-Strasse 4, CH-8006 Zurich, Switzerland; [email protected]; [email protected]; [email protected]; [email protected] • Mena Schemm-Gregory, in memoriam • Jiří Frýda, Faculty of Envi-ronmental Science, Czech University of Life Sciences, Kamýcká 129, 165 21 Praha 6 – Suchdol, Czech Republic;[email protected] • Björn Kröger, Freie Universität Berlin, Geologische Wissenschaften, FachrichtungPaläontologie, Malteserstrasse 74-100, D-12249 Berlin, Germany; [email protected] • Paul D. Taylor, Department ofEarth Sciences, Natural History Museum, Cromwell Road, SW7 5BD London, United Kingdom; [email protected] •Mark A. Wilson, Department of Geology, College of Wooster, Wooster, OH 44691-2363; [email protected]

The eastern Anti-Atlas in Morocco is famous for its highlyfossiliferous Palaeozoic rocks. Many palaeontological andstratigraphical studies on the Devonian of the Tafilalt andthe Maïder have been carried out during the twentiethand twenty-first century (Clariond 1934a, b; Roch 1934;Termier & Termier 1950; Massa et al. 1965; Hollard 1967,1974, 1981; Alberti 1980, 1981; Becker & House 1994,2000; Bultynck & Walliser 2000a, b; Klug 2001, 2002;Klug et al. 2008a, b; De Baets et al. 2010; Franchi et al.2012). Well-exposed sedimentary successions of the Early

Devonian age crop out in the Tafilalt, e.g. between the Je-bel Ouaoufilal and the mine of Filon 12 (Fig. 1). Thesemore or less complete exposures, the local abundance offossils and their sometimes excellent preservation yield thepossibility to study Early Devonian faunal associations intheir stratigraphic context in the Tafilalt.

In this study, we describe and figure well-preservedmacrofossils of five Early Devonian faunules (earliestLochkovian to early Emsian, according to Bultynck &Walliser 2000b), which comprise a highly diverse Pragian

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fauna of the Jebel Ouaoufilal (Filon 12). We found speciesof cephalopods, gastropods, anthozoans, brachiopods,trilobites, bivalves, crinoids, hederelloids, rostroconchs,hyolithids and machaerids, the larger representatives ofwhich are sometimes covered by epizoans. Our main focusis on these Early Devonian faunules, which we examinedfor alpha diversity and palaeoecology. Fluctuations in spe-cies richness and palaeoecology during the Early Devonianof Morocco have been noted before by several authors(e.g., Massa et al. 1965, Hollard 1974, Belka et al. 1999,Klug et al. 2008a, Kröger 2008). Belka et al. (1999) andKröger (2008) recognized an increase in diversity, espe-cially in benthic species, over time during the Early Devo-nian of the Tafilalt and correlated it to a change in oxygencontent at the sea floor. This hypothesis was, however, notyet tested by collecting and analysing quantitative data.Kröger (2008) examined species richness and compositionof the cephalopod associations at the Filon 12-section, butstatistical analyses on the species diversity of all groups ofmacroscopic invertebrates over the whole Early Devonianinterval have not been executed before. A palaeoecologicalstudy of the Tafilalt including two Emsian faunules waspublished by Klug et al. (2008a). They observed an ecolog-ical change in the early Emsian (Zlíchovian, sensu Bul-tynck & Walliser 2000b) based on two species richfaunules that differ in species composition.

To investigate the changes in species richness and thelocal palaeoecology from the earliest Lochkovian to theearly Emsian in the Tafilalt quantitatively, we analysedthe faunules based on alpha diversity and ecospace use.The term alpha diversity was introduced by Whittaker

(1960, 1972), who described the species richness within asingle habitat or community. The three-dimensional dis-play of ecospace utilisation, developed by Bush et al.(2007), consists of multiple ecological parameters, towhich each species can be assigned. These parameters arevertical tiering, motility level and feeding mechanism. Weemployed the same ecological parameters in our analysis.Additionally, the analysis of ecospace utilisation of theMoroccan Early Devonian faunules may reveal localmacroecological changes, which were linked with the “De-vonian nekton revolution” (Klug et al. 2010). The “Devon-ian nekton revolution” refers to a radiation of marinenektonic animals due to the rapid occupation of the watercolumn in the Devonian and the synchronous decrease inplanktonic and demersal taxa.

The aims of our study are (1) to describe and illus-trate the well preserved macrofossil material of five suc-cessive Early Devonian faunules from the Tafilalt, (2) todescribe new taxa of the highly diverse Pragian faunule,(3) to quantitatively analyse changes in alpha diversityand ecospace use through the Early Devonian of thesouthern Tafilalt, and (4) to discuss possible environ-mental and evolutionary factors, which might have initi-ated, controlled or influenced these changes in diversityand ecology.

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All the fossil specimens listed, figured and described in thiswork come from the locality Jebel Ouaoufilal in the Tafilalt

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�� !" Field photograph of the outcrop near Jebel Ouaoufilal and Filon 12. Image courtesy of Richard Hofmann (Zurich).

of Morocco (e.g. Klug et al. 2000, 2013). This locality issometimes also called Filon 12 (e.g., Kröger 2008) after thelocal mine, which is world-renowned for its excellent va-nadinite crystals and associated minerals.

The associations described here are earliest Lochkov-ian to early Emsian age (“Scyphocrinites”/Camarocrinuslevel to Erbenoceras/Anetoceras level; see Bultynck &Walliser 2000b, Corriga et al. 2013). They were collectedfrom the extensive outcrops of Devonian strata between thesouthwestern flank of Jebel Ouaoufilal, which is composedof Early Carboniferous siliciclastics, and the mentionedmine Filon 12. This area is located 7 km W of Taouz and40 km S of the town of Rissani. The Early Devonian sec-tion can be found approximately at the coordinates N30°56´57˝, W 04°02´39˝.

The samples of early to late Lochkovian age were takenbed by bed, whereas the samples of the lowermostLochkovian, the Pragian and the early Emsian (early Zlí-chovian) strata were collected from clayey to marly inter-vals. These intervals lie between limestone beds that formshallow ridges (as visible in Fig. 1) and so, the transport ofmaterial was limited to a few meters. Therefore, the ac-cording data do probably contain a slight bias from time-averaging and the sampling method. Some of the fossils,such as the large loboliths of the crinoid Camarocrinus,were identified and counted in the field in order to keep theweight of samples reasonably low. The remaining, mostlysmall, specimens of the faunules were quantitatively sam-pled and studied in the lab for species identification andsubsequent specimen counts (abundance). We definedwhole fossils or fossil fragments as single individuals ex-cept fin spines of the acanthodian genus Machaeracanthus.Machaeracanthus possessed four fin spines (Südkamp &Burrow 2007), and in order to account for the possibilitythat some of the fragments might have been parts of onespine or of several spines of one individual, we divided thetotal number of spines by two (although this number is ad-mittedly arbitrary). Similarly, we did not include crinoidcolumnals in the study, because it was impossible to esti-mate the number of crinoid individuals since the crinoidswere never entirely articulated. In the case of the genusCamarocrinus, we counted the loboliths, because only oneof these buoyancy devices was present in each individual(Haude 1992).

The species counts were plotted as histograms perfaunule and per ecological guild per faunule. Abundanceand diversity data were rarefied for each faunule with thesoftware package PAST (Hammer et al. 2001), in order toreduce the sampling bias and to compare more objectivelythe species richness of the faunules, which are of differentsample sizes (Sanders 1968).

After these investigations, we grouped all taxa accord-ing to ecological categories compiled by Bush et al. (2007),who classified marine animals with respect to vertical tier-

ing, motility level and feeding mechanism. Vertical tieringis defined by the vertical distribution of animals in or on thesediment as well as in the water column and includes thecategories pelagic, erect, surficial, semi-infaunal, shallowinfaunal und deep infaunal (Ausich & Bottjer 1982, Bottjer& Ausich 1986, Bush et al. 2007).

To the pelagic group, we assigned animals that lived inthe water column. This includes the nekton such as fishesand certain cephalopods and the plankton such as the spe-cialized genus Camarocrinus, which probably drifted atthe water surface with its lobolith functioning as a buoy(Haude 1992). All other crinoid species were stalked andthus were interpreted as erect benthonic animals, whichwere attached to the seafloor or to objects on the seafloorwith their body parts reaching more or less far into the watercolumn. Less highly erected benthonic organisms such asbrachiopods, bryozoans, corals, gastropods, hyoliths, he-derelloids, phyllocarids, trilobites, and some bivalve spe-cies, we allocated to the category surficial. Rostroconchs,pyrgocystid edrioasteroids and some bivalves belonged tosemi-infaunal animals (Kříž 2000, Klug et al. 2008a).Machaerids, the bivalve Panenka and palaeotaxodont bi-valves are assigned to the shallow infaunal benthos,whereas only one deep infaunal bivalve species was found(Kříž 2000, Klug et al. 2008a).

Motility level refers to the locomotory capabilities oforganisms and is subdivided in six categories: freely, fastor slow motile animals, attached or unattached faculta-tively motile animals and attached or unattachednon-motile animals (Bush et al. 2007). Freely, fast motileorganisms are represented by fishes and some arthropodsbecause of their active locomotion by fins or legs in thewater column or on the sea floor. Cephalopods includingActinocerida, Pseudorthocerida, Oncocerida, and Ortho-cerida were either slow swimming or vertically migratinganimals that lived in the water column; they were ratherslow swimming compared to many modern decabrachiancoleoid cephalopods (e.g., Westermann 1999, Westermann& Tsujita 1999). Therefore, we assigned cephalopods to-gether with gastropods and machaerids to the freely mov-ing, slow motile category. According to Kříž (2000), mostbivalves included in the study are considered as unat-tached, facultatively motile animals or, in the case ofepibyssate bivalves, to attached, facultatively motile or-ganisms. Corals, crinoids, bryozoans, hederelloids, edrio-asteroids and brachiopods belonged to the non-motile/ at-tached animals (Racheboeuf 1990, Williams et al. 2000).The unattached non-motile category includes rostro-conchs, hyoliths and a species of brachiopods (?Dagna-chonetes sp.) (Williams et al. 2000).

“Feeding mechanism” categorises the behaviour of or-ganisms to acquire food and is grouped in six subdivisions:suspension feeding, surface deposit feeding, mining, graz-ing, predatory and other (Bush et al. 2007). We consider

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tabulate corals, brachiopods, rostroconchs, bryozoans, he-derelloids, crinoids, edrioasteroids, hyoliths, phyllocaridsand epibyssate bivalves as suspension feeders that fed onsmall planktonic organisms from the water column. Themajority of the bivalve species found in our Early Devon-ian faunules fed on buried nutrition and, therefore, miningwas the common feeding mechanism. The predatory cate-gory includes fishes, rugose corals, nautiloids and trilobitesthat were able to catch prey capable of escaping and resist-ing. The trilobites of the faunules include members of theOrders Phacopida and Corynexochida that were suppos-edly microphagous predators because of the shape of theircephalon, the way of the attachment of hypostomes and thewell-developed sensory fields (Eldredge 1971, Miller1976, Whittington 1988, Fortey & Owens 1999). We as-signed the cephalopod species to microphagous predatorsas well based on the actualistic knowledge about the diet ofmodern Nautilus species. However, no undisputablecephalopod beaks older than Late Devonian are known. Inthe crop and stomach contents of recent Nautilida, frag-ments of small crustaceans and molts of larger crustaceanswere found, which were detected by their chemosensorytentacles (Saunders & Ward 1987). Bush et al. (2007)grouped all remaining feeding behaviours together as“others”. We added the new category “coprophagy”. Two

coprophagous species in the study belong to a single gas-tropod Family, the Platyceratidae (Peel 1984; Gahn &Baumiller 2003, 2006; Webster & Donovan 2012; Dono-van & Webster 2013).

The classification of taxa according to vertical tiering,motility and feeding mechanism led to different combina-tions of ecological categories, which represent differentmodes of life (Bush et al. 2007). We evaluated how manyspecies and individuals per mode of life were present ineach faunule and how their relative abundances changedthrough time. Additionally, the trophic nucleus conceptwas used to reveal which species and modes of life domi-nate the faunules. The trophic nucleus is defined by thosespecies that contribute to 80% of all individuals of a fauna(Neyman 1967).

The morphometric parameters for the description ofcephalopod conchs were adopted from Teichert et al.(1964) and Kröger (2008). For the description of crinoids,we used terms and parameters adopted from Ubaghs(1978) and Jell & Jell (1999). The descriptive terms andmeasurements for the description of gastropods were de-rived from Cox (1960). Morphometric parameters of hede-relloids were adopted from Bassler (1953) and Taylor &Wilson (2008).

All the material used for this study is housed in thePaläontologisches Institut und Museum der UniversitätZürich (PIMUZ 30587–30662). A second sample of thenewly introduced hederelloid genus Filihernodia was de-posited in the Department of Earth Science of the NationalHistory Museum in London (NHMUK PI BZ 7494).

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The Moroccan Anti-Atlas is a broad NE-SW oriented Va-riscan anticlinorium, which makes up a part of the northernmargin of the Sahara Craton (Piqué & Michard 1989). Pa-laeozoic sedimentary rocks can be found along a NE-SWtrending axis in the Anti-Atlas and include Devonian out-crops that cover an area of about 20.000 km2 (Kaufmann1998; see Fig. 2A). Devonian rocks are exposed along E-Wtrending synclines in the Tafilalt but parts of these rocksare interrupted by reverse faults with strike-slip compo-nents (Toto et al. 2008). Especially, the Silurian and EarlyDevonian claystone successions were deformed and/ or cutby faults, which led to incomplete exposures of the EarlyDevonian rock succession in the Tafilalt, especially inparts of the southern flanks of the synclines. Some excepti-ons are probably the southeastern limb of the so-calledAmessoui-syncline between the abandoned village El At-rous, the mine of Filon 12, Taouz, and the Jebel Ouaoufilal(Klug et al. 2013, see Fig. 2B).

In the Early Devonian, the Anti-Atlas was situated be-tween 40° to 60° south (Scotese 1997). Moderately cool

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�� %" Geological map of Morocco. • A – distribution of Palaeozoicrocks in the Anti-Atlas. • B – map of Devonian outcrops in the Tafilalt.

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�� '" Devonian section at Jebel Ouaoufilal (Filon 12). Modified after Klug et al. (2013). The section on the right has the same scale in horizontal andvertical direction (1:10,000).

water temperatures on the shelves of western and northernGondwana have been confirmed by oxygen isotope analy-ses by van Geldern et al. (2006, see also Lubeseder et al.2009). Early Devonian deposits display a rather uniform,mostly clayey to marly facies as well as moderate thicknesschanges in the eastern Anti-Atlas (Belka et al. 1997,Kaufmann 1998).

In the late Silurian and early Lochkovian (Early Devon-ian) successions, claystones are prevalent (Hollard 1981).These often dark coloured claystones (which are lightgreyish to pinkish when weathered) contain intercalatedmarlstone and limestone successions, which were formedby concentrated accumulations of planktonic crinoids (e.g.the scyphocrinoid limestone in which scyphocrinoid ossi-cles occur in rock-forming numbers), few orthoconiccephalopod shells (although some thin limestone interbedsare almost entirely composed of cephalopod shells) and oc-casionally bivalves as well as platyceratid gastropods(Haude 1992, Brachert et al. 1992, Kříž 2000, Lubeseder etal. 2009, Corriga et al. 2013). The limestones were depos-ited under low oxygen conditions below the storm wave-base and are characterized by massive beds with a maxi-mum thickness of approximately 1 m near Filon 12 (Kříž1998, Kröger 2008, see Fig. 3).

In the early Lochkovian, approx. 40 m above theScyphocrinites Beds, marls with nodules dominated by theorthoconic cephalopod Plagiostomoceras culter are fol-lowing near Filon 12 (Kröger 2008, Klug et al. 2013). Inthe late Lochkovian succession of dark claystones, somelayers of dark grey to almost black more or less nodularlimestones occur, which contain cephalopods, nowakiids,bivalves and a moderately low diversity in benthic species.Despite the presences of some benthic species, the facies ofthese limestones in combination with the predominance offreely motile cephalopods points at low oxygen conditionsin and on the sediment (Klug et al. 2013).

In the Pragian and Emsian, the carbonate content of thesediments is clearly higher than in the Lochkovian depos-its, which is reflected by the dominance of marls and nodu-lar limestones (Hollard 1981, Lubeseder et al. 2009).These limestones are much lighter in colour suggestinga lower content in organic material which points at moreoxygenated conditions at the sea floor (Klug et al. 2013). Inthe Filon 12 section, Pragian strata contain four limestonelayers, which are alternating with nodular marly lime-stones with macrofossils (Kröger 2008). Especially in theclaystone succession above the limestone “K3” of Kröger(2008), several nodular limestone layers occur, which arevery rich in macrofossils such as trilobites, cephalopods,echinoderms, gastropods, tabulate and rugose corals (Kluget al. 2008a, Kröger 2008). The last limestone bed is over-lain by light greyish marls, which are overlain by greenishclaystones with limonitised faunas as well as limonitepseudomorphoses after pyrite. The localisation of the

boundary between the Pragian and the Emsian is highlycontroversial and here, we were following the stratigraphyintroduced by Bultynck & Walliser (2000a, b) for theTafilalt. Accordingly, the Pragian-Emsian boundary isprobably located near the base of the greenish claystones inthe Tafilalt (Walliser 1984, 1985; Bultynck & Walliser2000b; Klug et al. 2008a; Klug et al. 2013; Kröger 2008).

In the greenish claystones, a highly fossiliferous faunaof the early Emsian, “faunule 1” sensu Klug et al. (2008a)and De Baets et al. (2010), occurs. These claystones arefollowed by the “Deiroceras limestone” with abundantlarge Deiroceras hollardi and are overlain by claystones of“faunule 2” described by Klug et al. (2008a) in the northernTafilalt (see also Kröger 2008). The subsequent Erbeno-ceras Limestone consists of fossiliferous wackestones con-taining dacryoconarids in great abundance as well as sub-ordinate crinoid ossicles, corals, trilobites, ostracods,bivalves, brachiopods, and the early ammonoids Erbeno-ceras advolvens as well as Anetoceras sp. (Klug 2001, DeBaets et al. 2013).

The Erbenoceras limestones are overlain by an over100 m thick claystone sequence. These claystones were de-posited during and after the initial Daleje transgression(see, e.g., Kröger 2008, Lubeseder et al. 2009, Klug et al.2013). During the late Emsian, the carbonate content in-creased more or less steadily and stayed high during muchof the Middle and parts of the Late Devonian succession inthe Amessoui Syncline (Döring 2002, Fröhlich 2004). Inthe late Middle Devonian, a platform and basin topographybegan to form, driven by early Variscan tectonic move-ments and differential subsidence (Wendt et al. 1984;Wendt 1985, 1988). Four depositional areas developed inthe Anti-Atlas: the Maïder Platform, the Maïder Basin, theTafilalt Platform and the Tafilalt Basin (Wendt 1988,Belka et al. 1997, Klug & Korn 2002).

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This faunule was found in the basalmost rocks of the sec-tion, representing the Silurian-Devonian boundary beds.This is corroborated by the presence of the plate-loboliths(“Platten-Lobolithen”) and conodonts (Haude & Walliser1998, Corriga et al. 2013). We found four taxa includingspecies of Camarocrinus, orthoconic cephalopods, bival-ves and platyceratid gastropods. Ecospace consists of fourmodes of life (see Fig. 5). The trophic nucleus includes Ca-marocrinus sp. and an orthocerid species, which belong tothe unattached pelagic, non-motile suspension feeders andpelagic, slow motile predators.

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The second faunule of the section is of early Lochkovianage (Belka et al. 1999). Species richness is limited to sixtaxa that include infaunal bivalves, a species each of cepha-lopods and of tabulate corals. Ecospace utilization is limi-ted to three modes of life (see Fig. 5). The trophic nucleuscontains the orthocerid species Plagiostomoceras culter,which is a pelagic, slow motile predator.

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This faunule of our Jebel Ouaoufilal section is of late Loch-kovian age (Klug et al. 2013) and species richness is com-posed of 20 species, which comprises bivalves, gastropods,orthocerids, rare brachiopods and one bryozoan species.The ecospace utilization includes six modes of life (seeFig. 5). Four species contribute to the trophic nucleus: Tem-peroceras sp. 1, Gastropoda ind. 6, Gastropoda ind. 1 and

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�� )" Alpha diversity of the species found in the earliest Lochkovian to early Emsian rocks at the Jebel Ouaoufilal (Filon 12) in the Tafilalt (Mo-rocco); dots above the bars mark the dominant species of the faunules. • A – Silurian-Devonian boundary, all 4 sampled taxa depicted. • B – earlyLochkovian, all 6 sampled taxa depicted. • C – late Lochkovian, only 10 taxa out of 20 sampled taxa illustrated. • D – Pragian, only 10 taxa out of 87 sam-pled taxa illustrated. • E – early Emsian, 10 taxa out of 42 sampled taxa illustrated. • F – overview of the changes in alpha diversity during the Early Devo-nian of the Taouz area. Additional taxa are listed in the appendix (Table 1).

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Actinopteria cf. decussata, which had modes of life such asfreely pelagic, slow motile predators, freely surficial, slowmotile grazers and attached semi-infaunal, facultativelymotile suspension feeders.

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The faunule was found in the Pragian rocks of the sectionat the Jebel Ouaufilal (Klug et al. 2013). The species rich-ness consists of 86 species and contains many orthoce-rids, actinocerids, oncocerids, gastropods, brachiopods,bivalves, crinoids, tabulate and rugose corals, arthropods,rostroconchs, hyolithids, machaerids and a new hederel-loid genus. 12 modes of life contribute to the ecospace ofthis faunule (see Fig. 5). The trophic nucleus comprises27 species and includes nautiloids, tabulate and rugosecorals, gastropods, brachiopods, bivalves and trilobites.These species had six different modes of life: freely pela-gic, slow motile predators, attached surficial, non-motilesuspension feeders, surficial, slow motile grazers, atta-ched surficial, non-motile predators, surficial, fast motilepredators, unattached shallow infaunal, facultatively mo-tile miners.

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The faunule belongs to the uppermost sediments of the sec-tion and is of early Emsian age (“faunule 1” of Klug et al.2010, 2013). The faunule is composed of 42 species ofbivalves, bactritids, orthocerids, actinocerids, oncocerids,gastropods, arthropods, corals, machaerids, fishes, and ed-rioasteroids. The ecospace consists of 14 modes of life (seeFig. 5). The trophic nucleus contains five species: Devono-bactrites obliquiseptatus, Phestia rostellata, Murchisoni-ceras murchisoni, Nuculoidea grandaeva, Orthotheca sp.These species belong to four modes of life such as freelypelagic, slow motile predators, unattached deep-infaunal,facultatively motile miners, unattached shallow-infaunal,facultatively motile miners and unattached surficial, non-motile suspension feeders.

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To investigate changes in alpha diversity and faunal com-position during the Early Devonian of the Taouz area, weidentified as many of the fossils as possible and counted alltaxa of every faunule and compared their abundances. At

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�� +" Changes in ecospace use during the Early Devonian (Silurian-Devonian boundaryto early Emsian) of the Taouz area.

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the Silurian-Devonian boundary, the species richness waslow. We found fossils of four species although we havemissed some species of orthocerids, which according toKröger (2008) occur therein. This is due to the fact thatsome layers are composed of almost only cephalopod shellswhile others are strongly dominated by Camarocrinus re-mains, i.e., when sampling for cephalopods, the focus is onthe cephalopod strata but we focused on the youngestcrinoid-layer. Nevertheless, we assume that the diversitywould be similarly low in the cephalopod layers and ben-thos is certainly also rare therein.

The diversity in groups and species increased slowlyfrom the early Lochkovian (6 species) to the late Loch-kovian (20 species). During the Pragian, the species diver-sity reached its Early Devonian maximum (86 species).The early Emsian faunule (42 species) had a lower speciesdiversity than the Pragian faunule.

These findings show that alpha diversity increased sig-nificantly from the Silurian-Devonian boundary to thePragian and decreased somewhat in the early Emsian(Fig. 4F). Rarefaction analyses indicated that in allfaunules except in the oldest (earliest Lochkovian) and theyoungest faunules (early Emsian) studied here further col-lecting would increase the number of species in the faunal

lists. The Pragian faunule still remains the most diverseamong the examined faunules and we probably sampledonly a fraction of the actual diversity (Fig. 6). In addition tothe species diversity, the taxonomic composition of the ex-amined faunules changed between the faunules and most ofthe species found in one faunule were not discovered infaunules of a different age. Furthermore, the number of in-vertebrate taxa of a higher systematic rank increasedthrough time and vertebrate remains appeared in the twoyoungest faunules.

��

Grouping of species according to the categories of tiering,motility and feeding mechanism resulted in different com-binations of categories reflecting different modes of life(Bush et al. 2007). These categories helped us investiga-ting changes in ecospace occupation during the Early De-vonian at our section of the Jebel Ouaoufilal (see Fig. 5).

The ecospace use at the Silurian-Devonian boundary ispoor in modes of life. Especially unattached pelagic ani-mals are common which are slow, facultative or non-motile and which fed on feces, small food particles in the

3%

�� ," Rarefaction analyses of all Early Devonian faunules (earliest Lochkovian to early Emsian) collected from the Jebel Ouaoufilal (Filon 12).

water column or small prey. Another mode of life is repre-sented by unattached shallow infaunal, facultativelymotile, mining bivalves. This result is, however, not en-tirely representative because of the relatively poor sam-pling. Most likely, we missed some species of orthoconesand possibly also bivalves.

In the early Lochkovian faunule, three modes of lifewere present of which only one (unattached shallowinfaunal, facultatively motile miners) dominated, reaching67% relative abundance. This mode of life is represented bybivalves, which show a slight increase in diversity comparedto the Silurian-Devonian boundary. Species of freely pe-lagic, slow motile predators that are represented by ortho-cones still exist but have a lower relative abundance (17%)than in the faunule of the earliest Lochkovian (25%). Withthe appearance of tabulate corals, a new mode of life (at-tached surficial, non-motile suspension feeders) occurred inthe sedimentary sequence at Filon 12, while two modes oflife from the Silurian-Devonian boundary (unattached pe-lagic, facultatively motile, coprophagous organisms or non-motile suspension feeders) apparently vanished with the dis-appearance of the early Lochkovian faunule.

In the late Lochkovian faunule, three additional modesof life appeared: freely surficial, slow motile grazers, at-tached surficial, facultatively motile suspension feedersand attached semi-infaunal, facultatively motile suspen-sion feeders. These modes were represented by gastropods,brachiopods, epibyssate and semi-infaunal bivalves. A to-tal of six modes of life existed then but none of them isclearly dominant. The highest relative abundances of theLochkovian faunule are represented by freely pelagic, slowmotile predators and freely surficial, slow motile grazers(26% for each).

In the Pragian faunule, six new modes of life appearedand so the number of modes of life had more than doubled.Especially surficial modes of life became more abundantand unattached semi-infaunal, non-motile and attachederect, non-motile suspension feeders were found for thefirst time. But of these 12 modes of life, the freely pelagic,slow motile predators have the highest relative abundanceof almost 34%. A further increase can be documented in“faunule 1” (Klug et al. 2008a, De Baets et al. 2010) of theearly Emsian (14 modes of life). No mode of life is signifi-cantly dominant but freely pelagic, slow motile predatorshave the highest relative abundance (21%). While five newmodes of life were present, we did not rediscover two thatexisted in the Pragian (attached erect and unattachedsemi-infaunal, non-motile suspension feeders). We foundfossils of freely pelagic, fast motile predators that were rep-resented by two fish species and a small increase in semi-infaunal and infaunal modes of life (attached semi-in-faunal, non-motile and attached shallow infaunal,facultatively motile suspension feeders, and unattacheddeep infaunal, facultatively motile miners).

We found evidence that the number of modes of life in-creased, and thus the ecospace use expanded from the Silu-rian-Devonian boundary towards the early Emsian. In theEarly Devonian of the Taouz area, we found only 19 of the216 theoretically possible modes of life. The number ofmodes of life recorded here corresponds well with the re-sults of Bush et al. (2007), who found 21 modes of life forthe mid Palaeozoic (Late Ordovician to Devonian) and 23to 25 for the late Cenozoic (Miocene to Pleistocene) ofNorth America and Europe. The specific types of modes oflife found in the Early Devonian of the Taouz area mostlycorrespond to those found in the mid Palaeozoic by Bush etal. (2007). Differences between the two studies (the typesof modes of life and their number) are influenced by thesize of samples and study area, the time span, preser-vational biases and different palaeoenvironments. For ex-ample, Bush et al. (2007) described a tropical environmentin the Palaeozoic of North-America while North Africawas situated in a more temperate climate (Kaufmann 1998,Scotese & McKerrow 1990).

As we examined all categories of ecospace occupationof Bush et al. (2007) separately, we found an increase ofecological categories that were occupied by species overtime in the section at the Jebel Ouaoufilal. The greatestchange occurred in the Pragian faunule. At this time, thenumber of surficial species, freely walking or slowlyswimming species and predator species had the highest ab-solute abundances within the classes of tiering, motilityand feeding mechanism (see Fig. 7).

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We examined changes in dominance based on analyses ofthe trophic nucleus. The trophic nucleus includes species,whose abundances contribute to 80% of the total numberof specimens per fauna (Neyman 1967). Moreover, we ana-lyzed to what modes of life (combination of tiering, moti-lity and feeding mechanism according to Bush et al. 2007)the dominant species belonged (see Table 1 in appendix).

At the Silurian-Devonian boundary, the trophic nucleuscontained two species and two different modes of life whileonly one species and a single mode of life were predomi-nant in the early Lochkovian faunule. Compared to this, thelate Lochkovian faunule contained more dominant speciesand modes of life (4 species, 3 modes of life). The distribu-tion of dominance changed profoundly from the Loch-kovian to the Pragian. 27 species and six modes of life werecontributing to the trophic nucleus of the Pragian faunule.A strong decrease in the number of predominant speciesoccurred in the early Emsian (5 species), whereas the num-ber of modes of life only decreased to four modes of life(see Fig. 4A–E).

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�� ." Changes in tiering, motility and feeding behaviour in the Early Devonian of the Taouz area. Ecological categories: Tiering – p: pelagic,er: erect, su: surficial, smi: semi-infaunal, si: shallow infaunal, di: deep infaunal; Motility – ff: freely fast, fs: freely slow, fu: facultatively unattached,fa: facultatively attached, nu: non-motile and unattached, na: non-motile and attached; Feeding type – s: suspension feeder, m: miner, g: grazer, pr: preda-tors, c: coprophagous (Bush et al. 2007).

The number of dominant species increased throughtime and the Pragian faunule with the highest species rich-ness also had the highest number of dominant species. Thetrophic nucleus includes species with pelagic modes of lifein the earliest Lochkovian and contains a lot of species ofdifferent groups and especially pelagic and surficial modesof life in the Pragian. In the early Emsian, the number ofdominant species strongly decreased, and pelagic, infaunaland surficial modes of life are dominating.

��

The alpha diversity increased from the Silurian-Devonianboundary to the Pragian and slightly decreased in the earlyEmsian of the Tafilalt. We found an expansion of the eco-space use and major changes in dominant species and modesof life in the trophic nuclei of the Early Devonian. Suchchanges in diversity and ecospace use can be caused by sealevel changes, changes in oxygenation of the sediment andwater on the sea floor, differences in salinity, or sedimentinput. A sampling bias might alter the primary data in suchway, that a trend can be seen, which does not correspond tothe actual original change in ecology. Sampling is affectedby, e.g., different sample sizes, poor sampling and time-averaging. The sampling bias due to sample size was redu-ced by rarefying the abundance data of the faunules by thesoftware package PAST (Hammer et al. 2001). In how farthe dimensions of the sampled intervals produced a bias isdifficult to estimate. In our opinion, however, this changein diversity and ecology is real, because even in singlebeds, many more species and fossil groups can be seen incross sections and furthermore, the Pragian diversity hasbeen corroborated by others, who examined, e.g., Pragiantrilobites (e.g. Alberti 1969). Time-averaging refers to theaccumulation of a local community over time, which doesnot represent the local community structure at a single mo-ment, because due to their short life spans, communities ac-cumulate faster than sediments (Walker & Bambach

1971). Therefore, short-term events are usually not visiblein fossil assemblages. Indeed, the long-term trends as seenin the section at the Jebel Ouaoufilal reflect extreme chan-ges in species richness and composition and the faunuleswere obviously affected by various environmental factors.Fluctuations in sea level or bottom oxygenation are pos-sibly the reasons for changes in diversity and ecospace uti-lization. Littoral parts of the sea are mostly low in diversitybecause of high water currents and perturbations while theenvironment of the shelves is more stable and therefore hashigher species richness (Ziegler 1972). The realms of thedeep sea are poor in species because the supply of organicfood usually decreases with water depth.

Many facts support the hypothesis that a sea level falland additional oxygenation of the sea floor occurred duringthe Early Devonian in the Tafilalt. The faunule of the Silur-ian-Devonian boundary was poor in species and modes oflife. Especially modes of life including pelagic species,such as orthocones, Camarocrinus and gastropods attachedto Camarocrinus were abundant. The faunule was domi-nated by two species (Orthocerida ind. 1, Camarocrinussp.) with following modes of life: freely pelagic, unat-tached non-motile suspension feeders and freely pelagic,slow motile predators. The missing benthos (except theinfaunal bivalve Panenka obsequens) and the dominatingpelagic species indicate low oxygen conditions on the seafloor and in the sediment during the earliest Lochkovian.Haude (1992) and Haude & Walliser (1998) reported thatthe planktonic scyphocrinoids existed from the Late Silur-ian to the early Lochkovian when the deposition of organicmaterial and therefore of black shales prevailed. This factin combination with the dominance of other pelagic organ-isms (nautiloids and graptolites) and the commonly darksediments rich in pyrite and organic matter, reflect hypoxicto anoxic conditions and thus poor living conditions at thebottom (Haude 1992). Kříž (2000) described a communityof large infaunal bivalves (“Panenka Community”), whichco-occurred with Scyphocrinites and cephalopods in theearliest Lochkovian of the Tafilalt. Based on the large size,

32

�� /" Lochkovian bivalves, Pragian trilobites and brachiopods from the Jebel Ouaoufilal in the Tafilalt. • A – Odontochile cf. hausmanni(Brongniart, 1822); fragment of the pygidium, × 1; PIMUZ 30645. • B–D – Reedops cf. cephalotes hamalagdianus Alberti, 1983; dorsal view of thecephalon, lateral view of the whole specimen, dorsal view of the thorax, × 1; PIMUZ 30646. • E–G – Reedops bronni (Barrande, 1846); lateral view of thewhole specimen, dorsal view of thorax and pygidium, × 1; PIMUZ 30647. • H, I – Cheirurus (Crotalocephalus) sp.; dorsal and lateral view of thecephalon, × 2; PIMUZ 30648. • J–N – Desquamatia sp.; lateral, dorsal, posterior, ventral and anterior view of the valves, × 1; PIMUZ 30649.• O–S – ?Aulacella eifeliensis (Verneuil, 1850); lateral, dorsal, posterior, ventral and anterior view of the valves, × 2; PIMUZ 30650. • T–X – Brachipodagen. et. sp. indet.; lateral, dorsal, posterior, ventral and anterior view of the valves, × 2; PIMUZ 30651. • Y–AC – ?Protathyris sp.; lateral, dorsal, poste-rior, ventral and anterior view of the valves, × 2; PIMUZ 30652. • AD–AO – Cingulodermis sp.; AD–AH – lateral, dorsal, posterior, ventral and anteriorview of the valves, × 2; PIMUZ 30653; AI–AO – lateral, dorsal, posterior, ventral and anterior view of the valves, × 2; PIMUZ 30654. • AP – aff.Eoglossinotoechia sp.; ventral view, × 2; PIMUZ 30655. • AQ, AR – Panenka princeps Barrande, 1881; dorsal and lateral views, × 1; PIMUZ 30656.• AS – Panenka humilis Barrande, 1881; lateral view, × 1; PIMUZ 30657. • AT – Neklania cf. resecta Barrande, 1881; insight view of the left valve, × 1;PIMUZ 30658. • AU – Jahnia aff. conscripta (Barrande, 1881); lateral view, × 3; PIUMZ 30659. • AV, AW – Actinopteria cf. decussata Hall, 1884;AV – lateral view, × 2; PIMUZ 30660; AW – lateral view of two specimens, × 2; PIMUZ 30660. • AX – Mytilarca sp.; lateral view, × 1; PIMUZ 30661.• AY – Patrocardia tarda (Barrande, 1881); lateral view, × 1; PIMUZ 30662.

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the low species diversity (Panenka obsequens, Panenkaaff. bellula and Dualina sp.) and the preservation of the bi-valves, he assumed a soft bottom environment with lowoxygen conditions and low water movement at the seafloor. Unfortunately, we found only three individuals ofPanenka obsequens and we did not find the other two spe-cies of the “Panenka Community” in our section at theJebel Ouaoufilal (Filon 12). But the low diversity, therather large size of the bivalves and the accompanying pe-lagic fauna indicates similar living conditions at the seafloor as reported by Kříž (2000).

In the early Lochkovian, the species diversity washigher because of the increased diversity in infaunal bi-valves (Genera: Panenka, Neklania, Jahnia) and the ap-pearance of a tabulate coral species. The ecospace use ex-panded slightly but the faunule still has a low diversity ofmodes of life, which include only pelagic and infaunal cat-egories. A cephalopod species, which belonged to freelypelagic, slow motile predators (Plagiostomoceras culter),strongly dominated the faunule. These facts all togethertestify that the bottom water was still similarly low in oxy-gen as during the Late Silurian and earliest Lochkovian.

In the late Lochkovian, the increase in alpha diversityand the gentle increase in modes of life with the first appear-ance of surficial gastropods and brachiopods in the sectionindicated slightly improved living conditions at the seafloor. We noted that the faunule was no longer dominated bypelagic species but also comprised surficial, freely slowmoving grazers (two gastropod species) and attached,semi-infaunal, facultatively motile suspension feeders(Actinopteria cf. decussata), which also reflected changes inthe environment. The species composition of the bivalveswas similar to the “Panenka-Jahnia-Neklania Community”described by Kříž (2000). This community contained a highdiversity of infaunal bivalves and also epibyssate bivalvesthat indicate improved living conditions at the bottom due tohigher oxygen content. In our sample, we found infaunal bi-valves such as Panenka humilis, Neklania cf. obtusa andJahnia sp., epibyssate bivalves such as Mytilarca sp. andPatrocardia tarda and the semi-infaunal bivalve Actino-pteria cf. decussata (see Fig. 8). Unfortunately, a lot of taxaof the “Panenka-Jahnia-Neklania” community were miss-ing. But higher diversity in infaunal species and appearanceof semi-infaunal and epibyssate bivalves reflect a better ven-tilated sea floor and therefore better living conditions forbenthic species in the late Lochkovian.

Subsequently, the extreme increase in alpha diversityfrom 20 species in the late Lochkovian to 86 in the Pragian,the increase in modes of life including surficial, erect,semi-infaunal and infaunal organisms and a highly diversetrophic nucleus (27 species, especially including pelagicand surficial modes of life) reflect good living conditionsfor benthonic as well as demersal life. In particular, thedrastic increase in benthic species and the colour change of

the sediment towards light grey permits the suppositionthat a steady rise of oxygen content near the seafloor oc-curred in combination with a sea-level fall. This is in accor-dance with the Devonian sea level curve of Morocco pre-sented by Kaufmann (1998) and is linked to a conspicuoussediment colour change, which occurs in the entire Tafilalt(Alberti 1981, Belka et al. 1999, Bultynck & Walliser2000b, Kröger 2008). Kröger (2008) correlated the in-crease in benthic taxa and cephalopods species at the Filon12 with the “pesavis Bioevent” or “Lochkovian-PragianBoundary Event” (Chlupáč & Kukal 1988, 1996; Schön-laub 1996), which coincides with global changes in sedi-ment colours, benthic diversity and/ or taxonomic compo-sition at the Lochkovian-Pragian boundary (Alberti 1969,Schönlaub 1996, Talent & Yolkin 1987, Talent et al. 1993,Kříž 1998, Ferreti et al. 1999, Chlupáč et al. 2000,García-Alcalde et al. 2000). These facts in combinationwith the Devonian eustatic sea level curves (Johnson et al.1985, 1996) revealed a change of oxygen content of the seafloor caused by a potential global sea level fall.

The early Emsian fauna (“faunule 1” of Klug et al.2008a) is also quite diverse although it contains less taxa(42 species), whereas the number of modes of life in-creased (from 12 modes of life in the Pragian to 14 in theearly Emsian). The high abundance of surficial species ledto the assumption that oxygen conditions in and on the sed-iment were similarly high as during the Pragian. Based onthe faunal composition and the facies, Klug et al. (2008a)concluded that the environmental conditions of the earlyEmsian can be considered as favorable and reasonably welloxygenated; however, the commonly limonitic and for-merly pyritic preservation of this fauna shows that onlythe top part of the sediment was oxygenated while deeperlayers were probably anoxic. Analysis on the trophic nu-cleus of the early Emsian faunule shows an elementarychange in the number of dominant species. There was anextreme decrease in dominant species (from 27 species inthe Pragian to 5 in the early Emsian) and a slight decreasein modes of life (from 5 modes of life in the Pragian to 4 inthe early Emsian) contributing to the trophic nucleus.Modes of life including pelagic and infaunal species weredominant in the early Emsian faunule, whereas only onesurficial species was prevalent. This fact, in combinationwith the limonitised macrofossils and the occurrence ofgreenish claystones reflect a less well-oxygenated bottomthan in the Pragian. Additionally, we found a high numberof infaunal bivalve taxa belonging to the Palaeotaxodontain the early Emsian faunule. Recent representatives of thisgroup are known to tolerate moderately low oxygen condi-tions in the sediment. The decrease in bottom oxygenationcould be explained by a transgression during the earlyZlíchovian (Kröger 2008, Lubeseder et al. 2009). Thisearly Emsian change in sea level possibly correlates to thetransgressive cycle Ib sensu Johnson et al. (1996).

33

��

In the present work, the discussed sea level changesmainly represent long-term trends in the Early Devonian ofthe Taouz area but naturally, many short-term fluctuationsin sea level occurred in this time span as well. Alternationsof limestones and claystones in our section at the JebelOuaufilal and in other sections in the eastern (Belka et al.1999, Kröger 2008) and southwestern Anti-Atlas (Lu-beseder et al. 2009) indicate numerous regional sea levelfluctuations. Comparisons between our results and the sec-tions of previous authors show similar changes in speciesrichness and ecospace use during the Early Devonian in theTafilalt, but lateral facies changes and diachronicity ofboundaries are possible and should be examined in greaterdetail in the future.

In Early Devonian faunules of the Tafilalt, the ecologi-cal turnover of the “Devonian nekton revolution” (Klug etal. 2010) is predominantly reflected in the faunal changesat the transition from “faunule 1” to “faunule 2” (Zlí-chovian, early Emsian) of Klug et al. 2008a, where parts ofthe abundance of non-ammonoid cephalopods were re-placed by rather diverse early ammonoids (Klug et al.2008a). Pelagic predators were already abundant in the Si-lurian and stayed common until the early Emsian but theymainly consist of slow motile microphagous orthocerids,actinocerids, oncocerids and bactritids. Nektonic predatorssuch as the acanthodian Machaeracanthus were found insmall numbers in the early Emsian faunule of our section.Demersal and planktonic species stayed dominant in thePragian and early Emsian faunules in the Taouz area.

*��

We examined five Early Devonian faunules near the JebelOuaoufilal in the southern Tafilalt for alpha diversity andecospace utilization. All 3376 specimens were identified asfar as possible and assigned to different modes of life inclu-ding the ecological groups tiering, motility and feedingmechanism according to the method introduced by Bush etal. (2007). These analyses revealed a strong increase inspecies richness and an extension in ecospace utilisationduring the Early Devonian (earliest Lochkovian to earlyEmsian sensu Bultynck & Walliser 2000b) of the Taouzarea. The highest number of species has been found in thePragian and the expansion of ecospace utilization reachedits maximum in the early Emsian. Especially taxa with sur-ficial, slow motile or predatory modes of life became mostdiverse in the Pragian and decreased somewhat in the earlyEmsian. The number of dominant species contributing tothe trophic nucleus increased from the Silurian-Devonianboundary to the Pragian and extremely decreased in theearly Emsian. All these changes reflect favorable conditi-ons for life on the sea floor during the Pragian of the Tafi-lalt. Increasing benthic and demersal diversity in the Prag-

ian coincides with colour changes in sediments and faunalchanges in the whole Tafilalt and other parts of the world atthe Lochkovian-Pragian boundary (Alberti 1969, 1981;Talent & Yolkin 1987; Talent et al. 1993; Kříž 1998;Schönlaub 1996; Ferreti et al. 1999; Belka et al. 1999; Bul-tynck & Walliser 2000b; García-Alcalde et al. 2000; Chlu-páč et al. 2000; Kröger 2008). The ecological change cor-relates with a steady rise of oxygen content near the bottomin combination with a regional if not global regression.This is in accordance with existing Devonian sea level cur-ves of Morocco (Kaufmann 1998) and eustatic sea levelcurves of Johnson et al. (1985; see also Haq & Schutter2008). In the early Emsian faunule of our section in the Ta-filalt, we found a decrease in benthic species with a simul-taneous increase in infaunal palaeotaxodont bivalves,which are tolerant to low oxygen conditions. This fact incombination with limonitised fossils and the sedimentsconsisting of greenish claystone documents a less oxygeni-zed sea floor than in the Pragian. This decline in oxygencontent at the sea floor could have been triggered by atransgression during the early Zlíchovian (Kröger 2008,Lubeseder et al. 2009) that possibly correlates with thetransgressive cycle Ib sensu Johnson et al. (1996).

Our analysis of the earliest Lochkovian to early Emsianfaunules (according to Bultynck & Walliser 2000b) of theTaouz area did not show a macroecological change similarto the “Devonian nekton revolution” (Klug et al. 2010).Demersal and planktonic species were predominant andwere not significantly displaced by nektonic organismssuch as acanthodian and ammonoids through time. Thegeological range and time span studied here limit the com-parison of our results with the diversity changes that oc-curred during the “Devonian nekton revolution” (Klug etal. 2010); this is linked with the fact that faunule composi-tions were examined here on a regional level only andsome important changes might have occurred within theEmsian, perhaps between “faunule 1” and “faunule 2”(Zlíchovian, early Emsian) of Klug et al. (2008a). How-ever, the “faunule 2” of Klug et al. (2008a) in our sectiondid not yield many fossils and it reflects only geographi-cally limited changes in palaeoecology.

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Phylum Mollusca Linnaeus, 1758Class Amphigastropoda Simroth, 1906Family Bellerophontidae M’Coy, 1851Subfamily Cymbulariinae Horný, 1963

New genus aff. Coelocyclus Perner, 1903

Remarks. – Horný (1963) established the new SubfamilyCymbulariinae within the Family Bellerophontidae, inclu-

31

ding four genera: Cymbularia Koken, 1896, PtychosphaeraPerner, 1903, Prosoptychus Perner, 1903, and CoelocyclusPerner, 1903. However, since his work of 1963, no revisionof this widely distributed group has been published.

New genus aff. Coelocyclus sp. nov.Figure 9AC–AE

Material. – One shell (PIMUZ 30604) from the Pragian ofJebel Oaoufilal in the Tafilalt (Morocco), housed in the Palä-ontologisches Institut und Museum der Universität Zürich.

Remarks. – Only a single shell is available, which resem-bles species of the genus Coelocyclus Perner, 1903. How-ever, its very sharp circumbilical ridge and very low whorlprofile differ from the morphological range of Coelocyc-lus. These distinct shell features suggest a position close toCoelocyclus but probably in an independent genus.

Occurrence. – Only one specimen from Jebel Ouaoufilal(Filon 12) in the Tafilalt of Morocco is known.

Class Gastropoda Cuvier, 1797

Remarks. – Family-level classification of Bouchet et al.(2005) is used in the following sections.

Family ?Oriostomatidae Wenz, 1938

Genus Oriomphalus Horný, 1992a

Type species. – Oriomphalus puellarum Horný, 1992a;Pragian, Early Devonian; basal part of Loděnice Lime-stone, Praha Formation, Prague Basin, Czech Republic.

Remarks. – Horný (1992a) placed two Early Devonian spe-cies, Oriomphalus puellarum Horný, 1992a and O. supra-liratus (Rohr & Smith 1978), within the new genus Oriom-phalus. O. supraliratus, was described from theLochkovian beds of the Canadian Arctic Islands and was

originally placed within the genus Cyclonema (Cyclo-nema) Hall, 1852 by Rohr & Smith (1978). The type spe-cies Oriomphalus puellarum Horný, 1992a comes from thebasal part of the Loděnice Limestone (Praha Formation,Pragian, middle Early Devonian) of the Prague Basin. La-ter, Frýda & Manda (1997) reported the occurrence ofanother species, O. aff. O. puellarum, from the Monograp-tus uniformis graptolite Biozone (early Lochkovian). Thehigher systematic position of the genus Oriomphalus is stilluncertain because of the lack of data on its protoconch mor-phology. Judging from teleoconch morphology, it pro-bably belongs to the archaeogastropod lineage and close tothe genus Australonema Tassell, 1980.

Oriomphalus multiornatus sp. nov.Figure 9A–O

Holotype. – The holotype (PIMUZ 30598) is figured inFig. 9M–O and is housed in the Paläontologisches Institutund Museum der Universität Zürich.

Type horizon and locality. – Pragian, pireneae Zone; SehebEl Rhassel Group, Jebel Ouaoufilal (Filon 12), Tafilalt,Morocco.

Material. – Seven shell fragments (PIMUZ 30587, PIMUZ30594–30598) from the Jebel Ouaoufilal in the eastern Ta-filalt, Morocco.

Etymology. – Multiornatus, combination of multi (much)and ornatus (decorated).

Diagnosis. – The species of Oriomphalus is ornamentedwith several distinct spiral threads crossed by dense collab-ral threads at early shell whorls and a wide pleural angle.

Description. – Description is based on all specimens(PIMUZ 30587, PIMUZ 30594–30598). Species with asmall, turbiniform shell having at least seven whorls; whorlprofile slightly shouldered in early whorls and rounded inlater whorls; whorl profile between the sutures stronglyconvexly arched; sutures distinctly impressed; pleural

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�� @" Pragian gastropods from the Jebel Ouaoufilal in the eastern Tafilalt. • A–O – Oriomphalus multiornatus gen. et sp. nov.; A–C – apical, lateraland basal views of the shell, × 2; PIMUZ 30594. D, E – apical and apertural views, × 2; PIMUZ 30595. F–I – apical, apertural, lateral and basal views, × 2;PIMUZ 30596. J, K – apical and apertural views, × 2; PIMUZ 30597. M–O – apical, apertural and lateral views, × 2; PIMUZ 30598. P–U – Australonemasp. nov. P, R – apical, apertural and basal views, × 2; PIMUZ 30599. S–U – apical, apertural and basal views, × 2; PIMUZ 30600. V, W – ?Spirina sp.; dor-sal and lateral view, × 2; PIMUZ 30601. • X, Y – Palaeozygopleura sp. nov.: lateral and apertural views, × 1; PIMUZ 30602. • Z–AB – Orthonychia sp.;apertural, lateral, dorsal views, × 1; PIMUZ 30603. • AC–AE – new genus aff. Coelocyclus sp. nov.; apertural, lateral and dorsal views, × 1; PIMUZ30604. • AF–AH – aff. Tychobrahea Horný, 1992a; apical, lateral and basal views, × 2; PIMUZ 30605. • AI–AN – Rihamphalus gracilis (Říha, 1938);AI, AK – apical, lateral and basal views, × 1; PIMUZ 30606; AL–AN – apical, apertural and basal views, × 2; PIMUZ 30607. • AO, AP – Paraoehlertiasp.; apical and lateral views, × 3; PIMUZ 30608. • AQ–AT – Eohormotomina restisevoluta gen. et sp. nov.; AQ, AR – apical and lateral views, × 3;PIMUZ 30609; AS, AT – apical and lateral views, × 3; PIMUZ 30610. • AU, AV – Umbotropis sp.; apical and lateral views, × 1; PIMUZ 30611.

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angle is about 90o; early whorls slightly shouldered; adultwhorls rounded; shell base narrowly phaneromphalous.Shell ornamentation consists of several distinct spiralthreads, only on the upper and median portion of the whorlcrossed by collabral threads in early shell whorls. The spi-ral threads are crossed by more or less regularly spaced,dense, collabral threads, which are always weaker than thestronger spiral threads of young whorls; the spiral threadsdisappear on more adult whorls; circumbilical thread pre-sent on all whorls. The protoconch is unknown.

Remarks. – The shell of Oriomphalus multiornatus gen. etsp. nov. differs from all known species of the genus, suchas the Pragian O. puellarum and the Lochkovian O. supra-liratus as well as O. aff. puellarum, by a much wider pleu-ral angle. In addition, it has a more distinct shell ornamen-tation than O. puellarum and O. aff. puellarum; it is alsomore shouldered in early whorls (compare with Horný1992a, pl. I, figs 5–8 as well as Frýda & Manda 1997, pl. 8,figs 6, 7). The Lochkovian O. supraliratus has a muchhigher shell and less prominent collabral threads thanO. multiornatus gen. et sp. nov.

Occurrence. – The species is only known from the localityJebel Ouaoufilal in the eastern Tafilalt, Morocco.

Genus Australonema Tassel, 1980

Type species. – Cyclonema australis Etheridge, 1890; Prag-ian, Cave Hill Quarries, Lilydale Limestone Formation,Australia.

Remarks. – Tassell (1980) established the new genus Aus-tralonema for gastropods with numerous, closely spacedspiral elements of ornamentation and placed it within theSubfamily Gyronematinae Knight, 1956 of the Family Ho-lopeidae Wenz, 1938. Tassell (1980) described and trans-ferred six species from Early Silurian to Early Devonianstrata of Australia to the genus Australonema. He also sug-gested the replacement of the Wenlockian Cyclonema cari-natum var. multicarinatum Lindström, 1884 to Australo-nema. Later, Gubanov & Yochelson (1994) described anadditional new species of Australonema, namely A. varva-rae from the Wenlockian of Siberia. The same authors alsoquestioned the higher taxonomy of Australonema and no-ted a similarity of this genus to Oriostoma Munier-Chalmas, 1876. Gubanov & Yochelson (1994) pointed outthat the presence of a shallow but distinct umbilicus in Orio-stoma distinguishes Australonema from the latter genus.However, as shown by Tassell (1980), the shells of Austra-lonema species including the type species A. australis(Etheridge, 1890), also have a well developed umbilicus.Yochelson & Linsley (1972) described and illustrated a

specimen of Cyclonema lilydalensis Etheridge, 1891(= A. lilydalensis) with an in situ paucispiral operculum.A paucispiral operculum was also found in the WenlockianA. varvarae Gubanov & Yochelson, 1994 and by Horný(1998), in the Pragian A. cf. guillieri (Oehlert, 1881). Re-cently, two new species were described from the Praguebasin: A. blodgetti Frýda & Manda, 1997 from the Mono-graptus uniformis graptolite Biozone (early Lochkovian)and A. havliceki Frýda & Bandel, 1997, from the upper-most part of the Třebotov Limestone (Daleje-TřebotovFormation, lowermost Eifelian).

Australonema sp. nov.Figure 9P–U

Material. – Two shells (PIMUZ 30599–30600) from thePragian of the Jebel Ouaoufilal in the Tafilalt (Morocco),housed in the Paläontologisches Institut und Museum derUniversität Zürich.

Remarks. – The specimens of Australonema sp. nov.clearly belong to the genus Australonema. This species dif-fers from all known Devonian species of Australonema bya very low spire. However, the limited material and the pre-servation do not suffice to introduce a new species.

Occurrence. – Only two specimens have been found whichare limited to the southern Tafilalt in Morocco.

Family Eotomariidae Wenz, 1938

Genus Paraoehlertia Frýda, 1998a

Type species. – Paraoehlertia parva Frýda, 1998a; earliestEifelian, Middle Devonian; uppermost part of the TřebotovLimestone, Daleje-Třebotov Formation, Prague Basin,Czech Republic.

Remarks. – The genus Paraoehlertia differs from the clo-sely related genus Oehlertia in its wide periselenizonearea, minutely phaneromphalous base, and opisthoclineapertural margin below the selenizone. The genus Para-oehlertia unites several, hitherto undescribed species fromthe Lower Devonian of Europe. The archaeogastropod pro-toconch type (Frýda 1998a) in the type species places thisgenus within the Subclass Archaeogastropoda.

Paraoehlertia sp.Figure 9AO, AP

Material. – A single specimen (PIMUZ 30608) with nicely

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preserved ornamentation (PIMUZ X) from the Jebel Oua-oufilal in the eastern Tafilalt (Morocco).

Remarks. – Only a single shell is available. However, itswell-preserved ornamentation shows all diagnostic charac-ters of the genus Paraoehlertia. Ongoing revision of theEarly Devonian gastropods from Europe and Australia (byJiří Frýda) revealed several already described species ofParaoehlertia that were placed in different eotomariid ge-nera. Species-level placement of the Moroccan Paraoeh-lertia will be possible after finishing the ongoing revisions.

Family Gosseletinidae Wenz, 1938

Genus Umbotropis Perner, 1903

Type species. – Umbotropis albicans Perner, 1903; LowerDevonian; Prague Basin, Czech Republic.

Remarks. – The type species Umbotropis albicans Perner,1903 comes from the Lower Devonian of the Prague Basin.Knight (1941) designated the holotype as derived from thespecimens shown by Perner (1903) in figs 6, 7 on pl. 42from the neighbourhood of the village Měňany. The typehorizon of this species is not clearly established but it pro-bably comes from limestones of Pragian age. Another Prag-ian species of the genus U. rihai Frýda & Manda, 1997,was described from the Prague basin. Besides these spe-cies, the Emsian U. mesoni Tassell, 1982 from the “Recep-taculites” Limestone of Australia is the only further speciesof this genus.

Umbotropis sp.Figure 9AU, AV

Material. – Two shells (PIMUZ 30611) from the Jebel Oua-oufilal in the Tafilalt (Morocco).

Remarks. – Both shells from the Jebel Ouaoufilal in theTafilalt clearly display generic features of the genus Umbo-tropis. The lack of additional shell characters (because oftheir weathered surface) does not allow their determinationon the species level.

Family Murchisonidae Koken, 1896

Genus Eohormotomina gen. nov.

Type species. – Eohormotomina restisevoluta gen. et sp.nov.; Pragian, Early Devonian; Jebel Ouaoufilal (Filon 12),Tafilalt, Morocco.

Etymology. – Eohormotomina, using the combination ofEos (= dawn) and the generic name Hormotomina.

Diagnosis. – A high-spired murchisoniid with selenizonebearing a medial spiral cord and bordered by two spiralcords. Selenizone situated at midwhorl height; whorl sur-face above and below selenizone bears prominent collab-ral, strongly prosocline costae forming a deep sinus.

Remarks. – The genus Eohormotomina is very close to thegenus Hormotomina Grabau & Shimer, 1909, based onMurchisonia maia Hall, 1861, from the Columbus Limes-tone (Eifelian) of Ohio. In contrast to Hormotomina, theshell of the Pragian Eohormotomina restisevoluta gen. etsp. nov. is ornamented with distinct collabral costae andhas a wider pleural angle. The shell of Eohormotomina alsoresembles Parahormotomina sibertae Blodgett, Frýda &Racheboeuf, 1999, from the Kersadiou Formation (middleGivetian) in the vicinity of Brest (Brittany, northwesternFrance). Eohormotomina differs from Parahormotominaby the absence of two spiral cords, one just above, and theother slightly below the suture. By the absence of a distinctspiral cord below its selenizone and distinct collabral cos-tae, Parahormotomina differs from the genus BouskaspiraFrýda 1999, which is based on the Pragian species Aclisinafugitiva Perner, 1907.

Eohormotomina restisevoluta sp. nov.Figure 9AQ–AT

Holotype. – Shell fragment (PIMUZ 30609) is figured inFig. 9AQ, AR. Paläontologisches Institut und Museum derUniversität Zürich.


Material. – The holotype (PIMUZ 30609) and a secondshell fragment (PIMUZ 30610) were collected from the Je-bel Ouaoufilal in the Tafilalt.

Etymology. – From restis (Latin) – rope and evolutus (La-tin) – enrolled; referring to the superficial resemblance toan enrolled pile of rope.

Diagnosis. – See generic diagnosis.

Description. – Description is based on all specimens(PIMUZ 30609–30610). Specimens with high spired shellswith at least nine whorls; spiral angle about 30 degrees. Se-lenizone raised, forming whorl periphery, situated at mid-whorl height, whorl profile above selenizone distinctly

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convex; selenizone bordered by prominent spiral cords andbearing a prominent median cord; width of selenizoneabout one third of the distance between sutures; upper spi-ral cord of selenizone slightly above mid-whorl height;whorl profile below selenizone flat on spiral whorls, gentlyrounded on final whorl; whorl surface above and below se-lenizone bears prominent collabral, strongly prosoclinecostae forming a deep sinus. The initial part of shells is un-known.

Remarks. – Horný (1992b) noted the occurrence of a singleshell fragment from the Loděnice Limestone (Pragian) ofBohemia, which shows diagnostic features of Hormoto-mina Grabau & Shimer, 1909 and might represent the old-est evidence for this genus. However, the shell surface ofthis poorly preserved fragment bears traces of distinct pro-socline cords as in Eohormotomina restisevoluta gen. et sp.nov. Therefore, it is possible that the shell fragment fromthe Loděnice Limestone belongs to our newly describedgenus Eohormotomina.

Occurrences. – The species has been found in the Tafilalt(Morocco) and is suspected to occur in the Loděnice Lime-stone of the Barrandian area (Czech Republic).

Family ?Euomphalidae de Koninck, 1881

Genus Rihamphalus Frýda, 1998b

Type species. – Porcellia gracilis Říha, 1938; Pragian,Early Devonian; Prague Basin, Czech Republic.

Remarks. – Říha (1938) placed his newly described specieswithin the genus Porcellia Léveillé and noted its similarityto the Silurian species P. sinistrorsa Perner, 1903. How-ever, P. sinistrorsa has a sinistrally coiled shell with a dis-tinct selenizone in contrast to Rihamphalus gracilis (Říha,1938). Frýda (1997) placed P. sinistrorsa within the newgenus Pernericirrus of the Family Porcelliidae, whichunites shells with dextral coiling in the early whorls andsinistral or planispiral coiling in the later whorls of the teleo-conch (Frýda & Blodgett 1998). The shell of R. gracilis(Říha, 1938) bears characteristic features of membersof the Family Euomphalidae. Nevertheless, diagnostic fea-tures of the Euomphalidae, such as the shape of the proto-conch, were not hitherto documented in Rihamphalus, andso its higher taxonomic position is uncertain.

Rihamphalus gracilis (Říha, 1938)Figure 9AI–AN

Material. – Five shell fragments (PIMUZ 30588, PIMUZ

30606–30607) were collected from the Jebel Ouaoufilal inthe eastern Tafilalt of Morocco.

Remarks. – All shell features of the shells coming from thelocality Jebel Ouaoufilal fit well to those of the type speciesRihamphalus gracilis. The genus was previously onlyknown from the Prague basin. Ongoing revision of the EarlyDevonian gastropods of Australia (by Jiří Frýda) revealedanother undescribed species of the genus Rihamphalus.

Occurrence. – The species is known from the Prague Basinin the Czech Republic and the Tafilalt in Morocco.

Superfamily Loxonematoidea Koken, 1889Family Palaeozygopleuridae Horný, 1955

Genus Palaeozygopleura Horný, 1955

Type species. – Zygopleura alinae Perner, 1907; Pragian,Early Devonian; Dvorce-Prokop Limestone, Praha Forma-tion, Prague Basin, Czech Republic.

Remarks. – Devonian species of this genus were recordedfrom Europe, North America, Australia, and Africa. Blod-gett et al. (1988, 1990) showed an Old World Realm distri-bution of Palaeozygopleura, which most probably arose inthe Rhenish - Bohemian Region of the Old World Realm,where the majority of the Early Devonian species and alsothe oldest species of this genus occur (Frýda 1993, Frýda &Blodgett 2004, Frýda et al. in press). Ongoing revision ofthe Early Devonian gastropods from Europe and Australia(by Jiří Frýda) revealed about ten hitherto undescribed spe-cies of Palaeozygopleura.

Palaeozygopleura sp. nov.Figure 9X, Y

Material. – 14 shell fragments (PIMUZ 30589, PIMUZ30602) from the Pragian of the Jebel Ouaoufilal in the Tafi-lalt (Morocco). The specimens are housed in the Paläonto-logisches Institut und Museum der Universität Zürich.

Remarks. – The fragments found between the Jebel Ouaou-filal and Filon 12 bear distinct shell characters, which allowsplacing them doubtlessly to the genus Palaeozygopleura.Judging from the fragment size, this species is probably thebiggest of all Palaeozygopleura species and by this charac-ter, it resembles a hitherto undescribed species from the Prag-ian of France. However, the shape of its asymmetric archedcostae (Fig. 9X, Y) differs from the species found in France.The Moroccan specimens belong to a new species of Palaeo-zygopleura, but incomplete shells (i.e., fragments formed

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only by two whorls) prevent to establish a new species. Be-side this new Moroccan species, the only other African spe-cies of Palaeozygopleura is P. vaneki Frýda, Ferrová, Ber-kyová, and Frýdová, 2008, reported by De Baets et al.(2010) from early Emsian strata of Morocco.

Occurrence. – The new species is restricted to the Tafilaltof Morocco.

Class Cephalopoda Cuvier, 1797Order Pseudorthocerida Barskov, 1963Family Spyroceratidae Shimizu & Obata, 1935

Genus Cancellspyroceras Kröger, 2008

Type species. – Orthoceras loricatum Barrande, 1868 by sub-sequent designation; Pragian Lower Devonian; lower part ofthe Dvorce-Prokop Limestone, Bohemia, Czech Republic.

Cancellspyroceras loricatum (Barrande, 1868)Figure 10A, B

Material. – Six fragments of phragmocones (PIMUZ30590, PIMUZ 30612) from the Jebel Ouaoufilal, Tafilalt,Morocco. PIMUZ 30612 with injury and with overgrowthof Filihernodia buccina Taylor & Wilson gen. et sp. nov.

Diagnosis. – See Kröger (2008).

Description. – The maximum observed length of the lar-gest fragment (PIMUZ 30612) is 264 mm. The widestconch diameter is 73.9 mm at the most abapical chamberand the smallest conch diameter is 30.5 mm at the mostadapical chamber. Slightly cyrtoconic shell, angle of ex-pansion approx. 11°. Ornamented shell with longitudinaland transverse lirae. Irregular distance between transverselirae, longitudinal lines and transverse lirae become narrowerfrom the abapical to the adapical part of the fragment. At adiameter of 49.4 mm, the ornamentation is interrupted by abig injury (height: 11.9 mm; width: 42.5 mm).

Occurrence. – Cancellspyroceras loricatum has been re-ported from the Pragian, Czech Republic and Morocco.

Family Pseudorthoceratidae Flower & Caster, 1935

Genus Geidoloceras Kröger, 2008

Type species. – Geidoloceras ouaoufilalense Kröger, 2008,original designation; Pragian, Early Devonian; Filon 12,Tafilalt, Morocco.

Geidoloceras ouaoufilalense Kröger, 2008Figure 11G, H

Holotype. – The specimen MB.C.9627 is housed in theMuseum für Naturkunde Berlin.

Type horizon and locality. – Pragian; bed KMO-II, Filon12 section, Tafilalt, Morocco.

Material. – 24 fragments of phragmocones (PIMUZ30591, PIMUZ 30616) from Jebel Ouaoufilal, Tafilalt,Morocco.

Diagnosis. – Emended after Kröger (2008): Orthoconic or

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�� !A" Large fragment of a Pragian nautiloid from the JebelOuaoufilal in the Tafilalt. • A, B – Cancellspyroceras loricatum(Barrande, 1868); lateral views of phragmocone fragment, × 0.5;PIMUZ 30612.

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slightly cyrtoconic shell with almost circular conchcross-section, angle of expansion approx. 10–12°. Shell or-namentation with transverse, narrow ridges. Siphunclesubcentral, expanding within the chambers, cyrtochoaniticseptal necks.

Description. – The length of the phragmocone fragmentPIMUZ 30616 is 31.5 mm with an orthoconic shell. Maxi-mum observed conch diameter 20.4 mm, minimum obser-ved conch diameter 16.4 mm, angle of expansion approx.12°. Shell ornamentation with oblique transverse narrowridges, 6 ridges per 1 mm. Three chambers are fully preser-ved; septal distance of the three abapical chambers is5.5 mm and 4.8 mm of the two adapical chambers. Siphun-cle subcentral; distance of the third septum (aboral) fromthe margin of the conch 9.3 mm on the dorsal side and7.9 mm on the ventral side. Siphuncle tube expands withinchambers, largest expansion of the connecting ring mea-sures 3.4 mm. Short septal necks cyrtochoantic, diameterof septal perforation 1.7 mm.

Occurrence. – The species is restricted to the Pragian in theTafilalt in Morocco.

Genus Subdoloceras Kröger, 2008

Type species. – Subdoloceras tafilaltense Kröger, 2008;Pragian, Early Devonian; Filon 12, Tafilalt, Morocco.

Subdoloceras atrouzense Kröger, 2008Figure 11I–L

Holotype. – Specimen MB.C.9595 in the Museum für Na-turkunde Berlin.


Material. – 46 phragmocones (PIMUZ 30592, PIMUZ30617–30618) from Jebel Ouaoufilal, Morocco, Prag-ian.


Description. – The length of the orthocone fragmentPIMUZ 30617 is 28.7 mm. Orthoconic conch, conicalshape, and circular conch cross-section. Maximum conchdiameter is 14.5 mm at the adoral site and minimum diame-ter is 6.3 mm at the aboral side, angle of expansion approx.15°. Shell ornamented with 10 fine transverse ridges, straight.Septal distance is 2.0 mm at the most adoral chamber and1.2 mm at the most aboral chamber. Siphuncle subcentraland thin.

Occurrence. – The species is restricted to the Tafilalt ofMorocco.

Order Orthoceratida Kuhn, 1940Family Orthoceratidae M’Coy, 1844

Tenuitheoceras gen. nov.

Type species. – Tenuitheoceras secretum gen. et sp. nov.,designated herein; Pragian, Early Devonian; Jebel Ouaou-filal (Filon 12), Tafilalt, Morocco.

Diagnosis. – Slender, orthoconic shell shape with com-pressed cross-section. Ornamented shell with faint obliquetransverse striae. Angle of expansion 6°, narrow septal dis-

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�� !!" Phragmocone fragments of Lochkovian and Pragian nautiloids from the Jebel Ouaoufilal in the Tafilalt. • A, B – Pseudendoplectoceraslahcani Kröger, 2008; lateral and septal views, fragments of the phragmocone, × 1; PIMUZ 30613. • C, D – Endoplectoceras sp.; lateral views,phragmocone with shell remains, × 2; PIMUZ 30614. • E, F – Tafilaltoceras adgoi Kröger, 2008; lateral and septal views, phragmocone incrusted bytabulate coral and crinoids, × 1; PIMUZ 30615. • G, H – Geidoloceras ouaoufilalense Kröger, 2008; lateral view and longitudinal section of an incom-plete phragmocone, × 1; PIMUZ 30616. • I–M – Subdoloceras atrouenze Kröger, 2008; I, J – lateral views of phragmocone fragment with epizoans,× 1.5; PIMUZ 30617; K–M – lateral and septal views of phragmocone with epizoans and machaerid fragment, × 1.5; PIMUZ 30618.• N, O – Subdoloceras engeseri Kröger, 2008; lateral views of phragmocone fragment, × 1; PIMUZ 30619. • P – Spyroceras cyrtopatronus Kröger,2008; lateral view of phragmocone fragment, × 1.5; PIMUZ 30620. • Q – Spyroceras patronus (Barrande, 1866); lateral view of phragmocone frag-ment, × 1.5; PIMUZ 30621. • R – Spyroceras latepatronus Kröger, 2008; lateral view of phragmocone fragment, × 1.5; NHMUK PI BZ 7494.• S–U – Arthrophyllum vermiculare (Termier & Termier, 1950); S – lateral view of phragmocone fragment, × 1.5; PIMUZ 30623; T – lateral view ofphragmocone with tabulate coral, × 1.5; PIMUZ 30624; U – longitudinal section of phragmocone, × 1; PIMUZ 30625. • V, W – Orthocyclocerastafilaltense Kröger, 2008; lateral and septal views of phragmocone, × 1.5; PIMUZ 30626. • X–Z – Tenuitheoceras secretum gen. et sp. nov.; longitudi-nal section and lateral views of phragmocone, × 1.5; PIMUZ 30627. • AA, AB – Adiagoceras sp.; lateral views of phragmocone, × 1; PIMUZ 30628.• AC, AD – Arionoceras kennethdebaetsi sp. nov.; longitudinal section and lateral view of phragmocone fragment, ×1; PIMUZ 30629.• AE, AF – Angeisonoceras reteornatum Kröger, 2008; lateral and septal view of phragmocone, × 1; PIMUZ 30630. • AG – Anaspyroceras sp.; lateralview of phragmocone fragment, × 1.5; PIMUZ 30631. • AH–AJ – Harrisoceras sp.; lateral and septal views of phragmocone fragment, × 1.5; PIMUZ30632. • AK – Plagiostomoceras culter (Barrande, 1866); lateral view of the mould of the body chamber and several phragmocone chambers, × 1;PIMUZ 30633. • AL–AM – Plagiostomoceras sp.; lateral views of phragmocone fragment, × 1.5; PIMUZ 30634. • AN–AO – Hemicosmorthocerassp.; lateral views of phragmocone fragment, × 1.5; PIMUZ 30635.

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tance approx. 0.3 of the conch cross-section. Siphunclesubcentral, long septal necks approx. 0.3 of the septal dis-tance, orthochoanitic. Diameter of septal perforation ap-prox. 0.1 of conch diameter.

Etymology. – After the similar genus Theoceras Kröger,2008 and tenuis (Latin) – narrow, referring to the compres-sed cross section.

Remarks. – The genus Tenuitheoceras is similar to Theoce-ras Kröger, 2008 in having a slender, orthoconic conch andnarrow septal distances. It differs in having a compressedconch cross-section, relatively long septal necks and a sub-central siphuncle instead of an eccentric siphuncle as inTheoceras. Moreover, the septal necks of the new genus donot form a funnel.

Included species. – Only the type species.

Tenuitheoceras secretum sp. nov.Figure 11X–Z

Holotype. – The specimen in Fig. 11X–Z, PIMUZ 30627,Paläontologisches Institut und Museum der UniversitätZürich.


Material. – Holotype: phragmocone fragment (PIMUZ30627) from Jebel Ouaoufilal (Filon 12), Tafilalt, Mo-rocco.

Etymology. – Secretum (Latin) – clandestine, secret, lurk-ing: referring to the fact that this species did not reveal itstraits superficially.

Diagnosis. – See generic diagnosis.

Description. – The holotype PIMUZ 30627 measures28.7 mm. Slender and orthoconic shape with compressedconch cross-section with the maximum diameter of13.3 mm, angle of expansion approx. 6°. Shell ornamentedwith 4 faint transverse striae per 1 mm. Three chambers arewell preserved; the maximum distance between two septais 4.3 mm in the most adapical chamber and the smallestdistance 3.9 mm in the second chamber. Siphuncle subcen-tral with distances of 5.8 mm dorsal and 4.9 mm ventralfrom the conch margin. Septal necks are orthochoanitic, re-latively long, length of the longest septal neck 1.4 mm inthe most adapical chamber. Diameter of septal perforationis 1.1 mm.

Occurrence. – Only the holotype from the Tafilalt of Mo-rocco is known.

Family Arionoceratidae Dzik, 1984

Genus Arionoceras Barskov, 1966

Type species. – Orthoceras canonicum Meneghini, 1857by subsequent designation; Wenlock, Silurian; Cea di SanAntonio, Fluminimaggiore, Sardinia, Italy.

Arionoceras kennethdebaetsi sp. nov.Figure 11AC, AD

Holotype. – The phragmocone fragment PIMUZ 30629(Fig. 11AC, AD), Paläontologisches Institut und Museumder Universität Zürich.


Material. – Only the holotype (PIMUZ 30629) from the Je-bel Ouaoufilal (Filon 12), Tafilalt, Morocco.

Etymology. – Species named after the Kenneth De Baets,honouring his contributions to research on Palaeozoic cep-halopods.

Diagnosis. – Orthoconic conchs with circular conchcross-section. Shell ornamentation with transverse striae isforming a deep sinus. Angle of expansion approx. 6°, sep-tal distance approx. 0.34 of the conch diameter. Siphunclecentral, short septal necks approx. 0.23 of the septal dis-tance, suborthochoanitic. Diameter of septal perforationapprox. 0.1 of conch cross-section, bigger than length ofseptal necks.

Description. – The maximum fragment length of the holo-type (PIMUZ 30629) is 27.5 mm. Orthoconic conch, circu-lar cross-section with the largest diameter 24.8 mm, angleof expansion approx. 6°. Fine transverse striae form a deepsinus, 4 striae per 1 mm, sutures straight. Three chambersare preserved, the highest septal distance measures 8.0 mmin the two most adapical chambers. Siphuncle central,small septal necks, suborthochoanitic, 1.8 mm long, septalperforation 2.2 mm in diameter.

Remarks. – Arionoceras kennethdebaetsi sp. nov. resem-bles A. capillosum Barrande, 1867 in having a similar angleof expansion, the same chamber/conch diameter ratio, thesame septal perforation/conch diameter ratio and septal

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neck morphology. But A. kennethdebaetsi sp. nov. differsfrom A. capillosum in having a much deeper sinus. The newlydescribed species has been found in the Pragian stage and isthe youngest Arionoceras species of Morocco so far.

Occurrence. – Only the holotype from the Tafilalt of Mo-rocco is known.

Family Geisonoceratidae Zhuravleva, 1959

Genus Angeisonoceras Kröger, 2008

Type species. – Orthoceras davidsoni Barrande, 1870; Up-per Silurian; Lochkov, Bohemia, Czech Republic.

Angeisonoceras reteornatum Kröger, 2008Figure 11AE, AF

Holotype. – The specimen MB.C.9946 is in the Museumfür Naturkunde Berlin.


Material. – 11 phragmocone fragments (PIMUZ 30593,PIMUZ 30630) from the Pragian, Jebel Ouaoufilal, Tafi-lalt, Morocco.


Description. – The phragmocone fragment PIMUZ 30630is 35.0 mm long, orthoconic, slightly acuminate, with a cir-cular conch cross-section. The maximum diameter at themost abapical part of the phragmocone is 19.9 mm; the mi-nimum diameter at the most adapical septum is 17.8 mm,angle of expansion approx. 5°. Ornamented shell with5 oblique transverse ridges per 1.0 mm, interrupted bysmooth longitudinal striae. Five chambers are preservedwith a measurable septal distance of 5.7 mm at the mostadapical chamber. Siphuncle subcentral, with a siphunclediameter of 2.0 mm at the adapical part of the fragment.

Occurrence. – The species is only known from the Tafilaltof Morocco.

Order Oncocerida Flower, in Flower & Kummel (1950)Family Nothoceratidae Fischer, 1882

Genus Tafilaltoceras Kröger, 2008

Type species. – Tafilaltoceras adgoi Kröger, 2008 by origi-

nal designation; Pragian, Early Devonian; Filon 12, Tafi-lalt, Morocco.

Tafilaltoceras adgoi Kröger, 2008Figure 11E, F

Holotype. – Specimen MB.C.9634, Museum für Natur-kunde Berlin.

Type horizon and locality. – Pragian; bed KMO-I, Filon 12section, Tafilalt, Morocco.

Material. – 1 phragmocone fragment (PIMUZ 30615)from Jebel Ouaoufilal, Tafilalt, Morocco.

Diagnosis. – Emended after Kröger (2008): Cyrtoconicshell expanding with slightly depressed conch cross-section; angle of expansion approx. 12 to 14°. Straightsepta; slightly concave. Siphuncle marginally, at the con-vex side of the shell; siphuncular diameter approx. 0.1 ofconch cross-section, suborthochoantic septal necks, dia-meter of septal perforation approx. 0.16 of conch diameter.

Description. – Phragmocone fragment PIMUZ 30615 is85.0 mm long with cyrtoconic shell and slightly oval conchcross-section. The maximum observed conch diametermeasures 39.4 mm and the smallest diameter 19.5 mm,angle of expansion is approx. 14°. Shell ornamentation isnot visible, because the specimen is completely encrustedby tabulate corals and a crinoid holdfast (Fig. 11E).20 chambers are preserved, which display straight sutures;the smallest septal distance at the most adapical chamber is2.9 mm and the largest septal distance at a diameter of30.8 mm. Siphuncle on the convex side of the conch, sip-huncle diameter 3.0 mm at the aboral side and 4.9 mm atthe adoral side.

Remarks. – The shell ornamentation of Tafilaltoceras ad-goi is still uncertain. The holotype found by Kröger (2008)and our new specimen (PIMUZ 30615) are extremely en-crusted by epizoans. The species is emended with regard tothe variability of the angle of expansion. In contrast to theholotype (angle of expansion: 12°), the new material reve-als an angle of expansion of 14°.

Occurrence. – Pragian, Tafilalt, Morocco.

Phylum Echinodermata Klein, 1734Class Crinoidea Miller, 1821Subclass Camerata Wachsmuth & Springer, 1885Order Monobathrida Moore & Laudon, 1943Suborder Compsocrinina Ubaghs, 1978

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Superfamily Hexacrinitacea Wachsmuth & Springer, 1885Family Hexacrinitidae Wachsmuth & Springer, 1885

Hexawacrinus gen. nov.

Type species. – Hexawacrinus claudiakurtae gen. et sp.nov., designated herein; Pragian, Early Devonian, JebelOuaoufilal (Filon 12), Tafilalt, Morocco.

Definition of genus. – Monocyclic calyx with conicalshape, smooth plates, 3 basals in the first row, 6 plates(5 radials and 1 anal plate?) in the second row. Length ofbasals approx. 0.6 of radial lengths, hexagonal. The radi-als are the biggest plates, heptagonal and hexagonal platesare alternating. The following primibrachs are smallerthan basals and radials, approx. 0.48 of the size of the ra-dials, hexagonal, higher in width than in length. The bran-ches are connected to the primibrachs, contact point onthe primibrachs is weakly developed. Two rows of inter-primibrachs, small, second row interprimibrachs are thesmallest plates, hexagonal. Sutures between plates arestrongly developed.

Remarks. – The new genus differs from HexacrinitesAustin & Austin, 1843 in lacking contact points for bran-ches at the anterior parts of the radials. The specimenshows some affinity to Wacrinus caseyensis Jell & Jell,1999. The sequence of the basals, radials, primibrachsand interprimibrachs is the same. Moreover, in both spe-cies, the contact points for the branching sclerites are situ-ated at the same position, namely at the anterior parts ofthe primibrachs. However, the proportions of the platesstrongly differ (e.g. basals are bigger and reach a higherlevel) and the plates are more symmetric than in Wacri-nus. The general calyx shape is rather conical than sub-spherical.


Hexawacrinus claudiakurtae sp. nov.Figure 12E–G

Holotype. – The specimen (PIMUZ 30636) is figured in

Fig. 12E, F and is housed in the Paläontologisches Institutund Museum der Universität Zürich.


Material. – The calyx fragment PIMUZ 30636 has beenfound at the Jebel Ouaoufilal in the eastern Tafilalt of Mo-rocco.

Etymology. – The species is named after the finder ClaudiaKurt (Basel), who kindly donated the specimen.

Diagnosis. – As for genus.

Description. – The calyx fragment PIMUZ 30636 is13.0 mm long and 12.6 mm wide and has a slightly conicalshape. All three basal plates are hexagonal and of similarshape and size. Their length is in between 3.6 mm and3.8 mm and reach approx. 0.6 of the length of the followingradial plates. Their maximum width is between 5.9 mm and6.1 mm. In the second row, just three plates are completelypreserved. They are the biggest plates of the calyx with si-milar size: length: 5.5 to 5.9 mm; width: 5.0 to 5.5 mm;they have an alternating heptagonal or hexagonal shape.All of them might be affiliated to radials. Two primibrachsare visible that are following two different radials. They are2.4 mm in length and 3.9 mm in width, symmetric and he-xagonal. They abut radials and interprimibrachs. One ofthe primibrachs shows a slight concave depression at theanterior margin and might be a potential contact point for asecundibrach. Two interprimibrachs on the first level arepresent. They are 2.9 to 3.3 mm long and 2.9 to 3.4 mmwide, hexagonal, slightly smaller than the primibrachs.Their margins abut radials, primibrachs and two interpri-mibrachs on the second level. Just two second row interpri-mibrachs are preserved. They are the smallest plates of thefragment, hexagonal and symmetric, 1.8 mm long and1.9 mm wide.

Occurrence. – A single specimen of Pragian age has beenfound at the Jebel Ouaoufilal in the eastern Tafilalt of Mo-rocco. Other occurrences are not known.

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�� !%" Hederelloids, crinoids, tabulate and rugose corals from the Pragian of the Jebel Ouaoufilal in the Tafilalt. • A–D – Filihernodia buccina gen.et sp. nov. Taylor & Wilson; A – large colony on a nautiloid, × 2; PIMUZ 30612; B–D – detail views of a small colony on a nautiloid, produced by a scan-ning electron microscope (SEM), B × 20 and C, D × 40; NHMUK PI BZ 7494. • E–G – Hexawacrinus claudiakurtae gen. et sp. nov.; lateral and basalviews of calyx fragment, × 2; PIMUZ 30636. • H – Tiaracrinus moravicus Ubaghs & Bouček, 1962; lateral view of the radial plate of the calyx, × 2;PIMUZ 30637. • I, J – Cleistopora cf. geometrica Milne-Edwards & Haime, 1851; view on three corallites and bottom, × 1; PIMUZ 30638.• K, L – Proclevia sp.; view on corallites and bottom side, × 1; PIMUZ 30639. • M–O – Favositidae Dana, 1846; M – lateral view, × 1; PIMUZ 30640;N, O – top and lateral view, × 1; PIMUZ 30641. • P, Q – unidentified tabulate coral; top and bottom view, × 1; PIMUZ 30642. • R – Aulopora sp.; smallcorallites on a hyolithid, × 2; PIMUZ 30643. • S – unidentified rugose coral; lateral view on a corallite, × 1.5; PIMUZ 30644.

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Subclass Disparida Moore & Laudon, 1943Order Cladida Moore & Laudon, 1943Suborder Compsocrinina Ubaghs, 1978Superfamily Belemnocrinacea S.A. Miller, 1892Family Zophocrinidae S.A. Miller, 1892

Genus Tiaracrinus Schultze, 1866

Type species. – Tiaracrinus quadrifrons Schultze, 1866;Eifelian, Middle Devonian; Freilingen Formation, Nollen-bach Member, Eifel, Germany.

Tiaracrinus moravicus Ubaghs & Bouček, 1962Figure 12H

Holotype. – Specimen BR 362, stored in the National Mu-seum of Prague, Czech Republic.

Type horizon and locality. – Emsian, Lower Devonian;Plumlov, Moravia, Czech Republic.

Material. – A cup fragment (PIMUZ 30637) collectedfrom the Jebel Ouaoufilal, Tafilalt, Morocco.

Diagnosis. – After Hauser (2008): Elongate conical calyx,with reduced, longitudinally oval radial rib field, 4 oval ra-dials with epispiral ridges and grooves and bulging three-part radial ring.

Description. – The maximum length of the fragmentPIMUZ 30637 is 7.8 mm and the maximum width is6.4 mm. The radial is large, has an ornamented surface dis-playing irregular ridges and bulges; its outline is rectangu-lar to slightly oval in shape, slightly convex with epispiralridges and grooves at the margins. There are 13 deep and12 shallow grooves alternating regularly on the right sideof the radial. On the left side, 14 deep grooves and 8 smal-ler grooves are visible. The radial becomes distally widerand displays 4 weak ridges.

Remarks. – In contrast to the type material, the here descri-bed new material, which consist of one radial plate, preser-ves the skeletal material and thus the surface ornamenta-tion. The assignment to Tiaracrinus moravicus is based onthe shape of the radial with the oval radial rib fields and thebroad radial channels. Between the radial rib fields, thesclerite’s surface displays an ornamentation of fine ribs, tu-bercles and nodes. In this respect, it resembles T. oehlerti,which has been suggested to be conspecific by Hauser(2008) and as sister species by Klug et al. (2013). Since thenew Moroccan material displays the morphology of theoutside of the radial, it can now be concluded that(1) T. moravicus and T. oehlerti are not conspecific and

should be kept as separate species, (2) the two species areindeed sister species, which (3) is now corroborated bythe coeval occurrence of the two species, which are clearlyolder than four other species of Tiaracrinus (T. rarus,T. jeanlemenni, T. tetraedra, T. quadrifrons). The here de-scribed new specimen represents the first record of T. mo-ravicus from Africa.

Occurrence. – An Emsian specimen has been found inPlumlov, Moravia, Czech Republic and a Pragian speci-men is here described from the Jebel Ouaoufilal, Tafilalt,Morocco.

Phylum Incertae sedisSuborder Hederelloidea Bassler, 1939

Remarks. – Hederelloids are among the most commonencrusting sclerobionts on Devonian hard substrates. Al-though traditionally considered as cyclostome bryozo-ans (e.g., Bassler 1953 in the bryozoan Treatise), theydiffer from cyclostomes and bryozoans in several impor-tant respects. Notably, hederelloid walls have a prisma-tic microstructure and the zooids are often larger thanthose found in any unequivocal bryozoans. Taylor &Wilson (2008) reconsidered the affinities of these colo-nial animals, noting similarities in zooidal budding pat-tern to that of the colonial phoronid worm Phoronis ova-lis. Despite the lack of a mineralized skeleton in recentphoronids, the resemblance led these authors to hypothe-size that hederelloids were an extinct (Silurian–Per-mian) clade of colonial phoronids that evolved calcifiedskeletons.

Family Reptariidae Simpson, 1897

Genus Filihernodia Taylor & Wilson gen. nov.

Type species. – Filihernodia buccina gen. et sp. nov. Tay-lor & Wilson; Pragian, Early Devonian; Jebel Ouaoufilal(Filon 12), Tafilalt, Morocco.

Etymology. – Alluding to its similarity with Hernodia Hall,1881, but having zooids that are very narrow proximally,hence fili from filum meaning “thread, string, filament,fibre”.

Genus definition. – Hederelloid with horn-shaped zooidsoriginating as narrow buds mid-way along the lengths ofparent zooids, alternately on the left and right sides, exceptat branch bifurcations, where a pair of daughter zooids ispresent, one on each side of the parent; new buds diverge atabout 60° from parent zooids and broaden distally.

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Remarks. – Five genera of hederelloids are currently recog-nized (Taylor & Wilson 2008). This new genus most clo-sely resembles Hernodia but differs in having zooids thatare very narrow proximally and broad distally, whereas thezooids of Hernodia have a uniform width. Indeed, the three-fold increase in width along the zooids of Filihernodia isan unusual attribute among hederelloids as a whole. Fur-thermore, slender zooids of Filihernodia rapidly separatefrom their parent zooids, giving the colony a more gracilemorphology than Hernodia in which daughter zooids di-verge at low angles and tend to adhere to the sides of theirparents for some distance before separating. The longitudi-nal ridges visible on the surfaces of some zooids of Filiher-nodia are lacking in other hederelloids, where transversegrowth bands tend to be better developed.


Filihernodia buccina sp. nov. Taylor & WilsonFigure 12A–D

Holotype. – The specimen (PIMUZ 30612) is figured inFig. 12A, and is housed in the Paläontologisches Institutund Museum der Universität Zürich.


Material. – Holotype: PIMUZ 30612 (large colony in Zu-rich). Paratype: NHMUK PI BZ 7494 (small specimenscanned at the NHM) in Fig. 12B, C. Both specimens havebeen found at the Jebel Ouaoufilal (Filon 12) in the Tafilaltof Morocco.

Etymology. – Buccina, L. for shepherd’s horn, in referenceto the shape of the zooids.

Diagnosis. – As for genus.

Description. – Colony (NHMUK PI BZ 7494) encrusting,uniserial, runner-like, comprising slender, gently sinuous,bifurcating branches formed of zooids budded approxima-tely mid-length on the side of a zooid from the precedinggeneration, alternately to the left and the right and diver-ging at up to 60°. Branch bifurcation is infrequent, arisingfrom budding of paired daughter zooids on the two oppo-site sides of a parent zooid, which is straighter than normal,the two branches diverging initially at about 90°. Early as-togeny is unknown.

Zooids tubular, horn-shaped, curved through an angleof up to about 45°, narrow proximally, gradually broaden-ing distally towards the terminal aperture. Total zooid

length 2.5–3.5 mm, proximal width about 0.12–0.14 mm,distal width 0.38–0.46 mm. Walls non-porous, marked byfaint transverse growth lines and up to three, low longitudi-nal ridges in some zooids. Aperture is subcircular andabout 0.26–0.28 mm in diameter.

Remarks. – As noted above, this new genus and speciesmost closely resembles Hernodia, which Bassler (1939)distinguished from other hederelloids on the basis of itselongate, club-shaped zooids budding alternately fromabout the middle of the sides of the preceding zooids. Bas-sler (1939) described eight species of Hernodia from theSilurian-Devonian of the United States, noting two furtherspecies from the Silurian and Devonian of Bohemia. Onenamed and one questionable species of Hernodia were sub-sequently described from the Devonian of Germany (Solle1952, 1968).

Compared to species of Hernodia, F. buccina has colo-nies of a more gracile appearance, with long, narrow zooidsthat diverge from their parent zooids at relatively high an-gles. Species such as H. tennesseensis Bassler, 1935 andH. concinna Bassler, 1935, by contrast, have low angles ofbud divergence, resulting in the new buds being alignedsubparallel and often initially adherent to the parent zooid.In terms of zooid size, F. buccina is most similar toH. davisi Bassler, 1935 from the Hamilton Group of theFalls of the Ohio, but zooids of this species maintain an al-most constant width along their lengths.

Occurrence. – The species is only known from the Pragianof the Jebel Ouaoufilal in the Tafilalt (Morocco).

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We greatly appreciate the support by the Swiss National ScienceFoundation (Project No. 200021–113956/1, 200020–25029, and200020–132870). Jens Koppka (Porrentruy) kindly checked ourdeterminations of the trilobites. Joachim Hauser (Bonn) helpedwith the determination of the new crinoid species. We thank thestudents of the University of Zurich who joined thepalaeontological fieldwork course 2012 in Morocco and helped tocollect the fossil material included in the study. Participation ofJiří Frýda was supported by the Grant Agency of the Czech Re-public (P210/12/2018).

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��!" Absolute abundances of the species found in the earliestLochkovian to early Emsian rocks at the Jebel Ouaoufilal in the easternTafilalt of Morocco, and their ecological classification after Bush et al.(2007). Ecological categories: Tiering – p: pelagic, er: erect, su: surficial,smi: semi-infaunal, si: shallow infaunal, di: deep infaunal; Motility – ff:freely fast, fs: freely slow, fu: facultative unattached, fa: facultative at-tached, nu: non-motile and unattached, na: non-motile and attached;Feeding mechanism – s: suspension feeder, m: miner, g: grazer, pr: preda-tors, c: coprophagous.

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?Class Filihernodia buccinagen. et sp. nov.

0 0 0 2 0 su, na, s

Stenolaemata Treptosomes orCytoporates

0 0 1 0 0 su, na, s

Crinoidea Hexawacrinusclaudiakurtae gen. etsp. nov.

0 0 0 1 0 er, na, s

Tiaracrinus cf.moravicus

0 0 0 1 0 er, na, s

Camarocrinus sp. 65 0 0 0 0 p, nu, s

Edrioasteroidea Rhenopyrgus flos 0 0 0 0 11 smi, na,s

Anthozoa Favositidae 1 0 0 0 19 2 su, na, s

Cleistopora cf.geometrica

0 0 0 4 0 su, na, s

Proclevia sp. 0 0 0 1 0 su, na, s

Tabulata 1 0 0 0 3 0 su, na, s

Tabulata 2 0 0 0 2 0 su, na, s

aff. Striatopora sp. 0 0 0 7 0 su, na, s

Auloporidae 1 0 3 0 0 0 su, na, s

Aulopora sp. 0 0 0 1 0 su, na, s

Rugosa 1 0 0 0 1 0 su, na, pr



Rugosa 4 0 0 0 17 22 su, na, pr




Trilobita Reedops bronni 0 0 0 1 0 su, ff, pr

Reedops cf.cephaloteshamalagdadianus

0 0 0 7 0 su, ff, pr

Pseudocryphaeus sp. 0 0 0 1 0 su, ff, pr

Pilletina zguidensis 0 0 0 0 60 su, ff, pr

Metacanthinawallacei

0 0 0 0 3 su, ff, pr

Odontochile cf.hausmanni

0 0 0 1 0 su, ff, pr

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Cheirurus(Crotalocephalus) sp.

0 0 0 2 0 su, ff, pr

Hollardops sp. 0 0 0 1 0 su, ff, pr

Paralejuruselayounensis

0 0 0 2 0 su, ff, pr

Paralejuruscampanifer

0 0 0 0 1 su, ff, pr

Malacostraca Nahecaris jannae 0 0 0 0 5 su, ff, s

Rhynchonellata Desquamatia sp. 0 0 0 4 0 su, na, s

?Aulacella eifeliensis 0 0 0 1 0 su, na, s

aff.Eoglossinotoechiasp.

0 0 0 1 0 su, na, s

Brachiopoda gen. etsp. indet.

0 0 0 6 0 su, na, s

?Protathyris sp. 0 0 2 9 9 su, na, s

Cingulodermis sp. 0 0 0 12 0 su, na, s

Arduspiriferarduennensis

0 0 2 0 1 su, na, s

Quadrithyristermierae

0 0 0 0 1 su, na, s

Strophomenata ?Dagnachonetes sp. 0 0 0 1 0 su, nu, s

Bivalvia Bivalvia ind. 1 0 0 10 0 0 ?

Panenka humilis 0 0 5 1 0 si, fu, m

Panenka hollardi 0 0 0 3 4 si, fu, m

Panenka princeps 0 1 0 0 0 si, fu, m

Panenka sp. 1 0 1 0 0 0 si, fu, m

Panenka obsequens 3 0 0 0 0 si, fu, m

Neklania cf. resecta 0 1 0 0 0 si, fu, m

Neklania cf. obtusa 0 0 1 0 0 si, fu, m

Jahnia sp. 0 0 1 0 0 si, fu, m

Jahnia aff.conscripta

0 5 0 0 0 si, fu, m

Patrocardia evolvensevolvens

0 0 0 0 1 su, fa, s

Patrocardiaexcellens sp.

0 0 0 0 10 su, fa, s

Patrocardia tarda 0 0 2 2 16 su, fa, s

Mytilarca cf.chemungensis

0 0 0 0 1 su, fa, s

Mytilarca sp. 0 0 2 0 0 su, fa, s

Actinopteria cf.decussata

0 0 11 0 0 smi, fa, s

Grammysioidea sp. 0 0 0 0 3 si, fa, s

Nuculoideagrandaeva

0 0 0 1 79 si, fu, m

Eonuculoma babini 0 0 0 0 2 si, fu, m

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Palaeoneiloemarginata

0 0 0 2 59 si, fu, m

Nuculites celticus 0 0 0 0 5 si, fu, m

Cucullella triquetra 0 0 0 0 52 si, fu, m

Phestia rostellata 0 0 0 0 149 di, fu, m

Rostroconchia Bohemicardiabohemicus

0 0 0 1 0 smi, nu,s

Hyolitha Orthotheca sp. 0 0 0 3 66 su, nu, s

Machaeridia Machaeridia 1 0 0 0 1 0 si, fs, m

Lepidocoleus rugatus 0 0 0 0 5 si, fs, m

Amphigastropoda aff. Coelocyclus sp.nov.

0 0 0 1 0 su, fs, g

Sinuitina sp. 0 0 0 0 5 su, fs, g

Crenistriella sp. 0 0 0 0 8 su, fs, g

Gastropoda Gastropoda ind. 1 0 0 20 0 0 su, fs, g

Gastropoda ind. 2 0 0 0 4 0 su, fs, g





Spirina sp. 0 0 0 1 0 su, fs, g

Oriomphalusmultiornatus gen. etsp. nov.

0 0 0 7 0 su, fs, g

Australonema sp.nov.

0 0 0 2 0 su, fs, g

Eotomarioidea ind. 1 0 0 0 1 0 su, fs, g

Eotomarioidea ind. 2 0 0 0 1 0 su, fs, g

Paraoehlertia sp. 0 0 0 1 0 su, fs, g

Umbotropis sp. 0 0 0 2 0 su, fs, g

Eohormotominarestisevoluta gen. etsp. nov.

0 0 0 2 0 su, fs, g

Rihamphalus gracilis 0 0 0 5 0 su, fs, g

aff. Tychobrahea sp. 0 0 0 1 0 su, fs, g

Loxonematoideaind. 1

0 0 0 7 0 su, fs, g

Loxonematoideaind. 2

0 0 0 3 0 su, fs, g

Palaeozygopleura sp.nov

0 0 1 14 0 su, fs, g

Palaeozygopleura sp. 0 0 0 0 34 su, fs, g

Pleurotomarioideaind., new genus?

0 0 0 2 0 su, fs, g

Lukesispira pulchra 0 0 0 0 9 su, fs, g

Planitrochus tardus 0 0 0 0 12 su, fs, g

Platyceratidae ind. 1 7 0 0 0 0 p, fu, c

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Orthonychia sp. 0 0 0 1 0 su, fs, c

Cephalopoda Orthocerida ind. 1 21 0 0 0 0 p, fs, pr

Orthocerida ind. 2 0 0 0 4 0 p, fs, pr




Endoplectoceras sp. 0 0 0 1 0 p, fs, pr

Pseudendoplectoceraslahcani

0 0 1 1 0 p, fs, pr

Bohemojovellania sp. 0 0 3 0 0 p, fs, pr

Tafilaltoceras adgoi 0 0 0 1 0 p, fs, pr

Pseudorthoceratidaeind. 1

0 0 1 0 0 p, fs, pr

Geidolocerasouaoufilalense

0 0 0 24 0 p, fs, pr

Subdolocerasatrouzense

0 0 0 46 0 p, fs, pr

Subdolocerasengeseri

0 0 0 40 0 p, fs, pr

Spyrocerascyrtopatronus

0 0 0 6 0 p, fs, pr

Spyroceraslatepatronus

0 0 0 1 0 p, fs, pr

Spyroceras patronus 0 0 0 3 0 p, fs, pr

Spyroceras sp. 0 0 0 0 9 p, fs, pr

Cancellspyrocerasloricatum

0 0 0 6 0 p, fs, pr

Arthrophyllumvermiculare

0 0 0 63 13 p, fs, pr

Murchisonicerasmurchisoni

0 0 0 0 96 p, fs, pr

Infundibulocerasbrevimira

0 0 0 0 18 p, fs, pr

Neocycloceras sp. 0 0 0 0 1 p, fs, pr

Michelinoceras sp. 0 0 0 1 0 p, fs, pr

Orthocyclocerastafilaltense

0 0 0 3 0 p, fs, pr

Orthocycloceras sp. 0 0 0 0 26 p, fs, pr

Tenuitheocerassecretum gen. et sp.nov.

0 0 0 1 0 p, fs, pr

Tibichoanoceras sp. 0 0 0 1 0 p, fs, pr

Arionoceraskennethdebaetsi sp.nov.

0 0 0 1 0 p, fs, pr

Adiagoceras sp. 0 0 0 2 0 p, fs, pr

Anaspyroceras sp. 0 0 0 1 0 p, fs, pr

Harrisoceras sp. 0 0 0 1 0 p, fs, pr

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Angeisonocerasreteornatum

0 0 0 11 0 p, fs, pr

Temperocerasaequinudum

0 0 0 3 0 p, fs, pr

Temperoceras sp. 1 0 0 122 0 0 p, fs, pr

Temperoceras sp. 2 0 0 0 0 36 p, fs, pr

Temperocerasludense

0 0 0 10 0 p, fs, pr

Sphaerorthoceratidae 0 0 0 1 0 p, fs, pr

Plagiostomocerasculter

0 121 0 0 0 p, fs, pr

Plagiostomocerashassichebbiense

0 0 0 0 7 p, fs, pr

Plagiostomoceras sp. 0 0 0 2 0 p, fs, pr

Hemicosmorthocerassp.

0 0 0 1 0 p, fs, pr

Bactritidae ind. 0 0 0 1 0 p, fs, pr

Devonobactritesobliquiseptatus

0 0 0 0 1604 p, fs, pr

Placodermi Brachythoracidplacoderm

0 0 0 0 5 p, ff, pr

Acanthodii Machaeracanthus cf.peracutus

0 0 0 0 18 p, ff, pr

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